loHlnn Untttrrattg
QlnlUgp nf Hib^ral ArtB
The Gift of . ..ir.ke TT^utWor.
BOSTON UNIVERSITY
GRADUATE SCHOOL
Dissertation
NEURO-MUSCULAR ACTIVITY IN THE
PEDAL WAVES OP HELIX
by
Blanche Brine Daly
(A.B,, Hunter College, 1913; M.Sc, Nev/ York University,
1915; M.A., Radcliffe College, 1928)
submitted in partial fulfilment of the
requirements for the degree of
Doctor of Philosophy
1935
4q ^ 11
-PhD
19 33
ACKNOWLEDGMENT
The writer wishes to express her
«
appreciation to Professor Brenton h. Lutz ,
Department of Biology, Boston University,
for his helpful advice and continued interest
in this work.
CONTENTS
CHAPTER I Introduction Page i
CHAPTER II Purpose of this investigation 5
CHAPTER III Review of the literature 10
CHAPTER IV Method and procedure 19
CHAPTER V Results and discussion 26
A. Creeping during vertical
ascension without loads 26
B. Comparison of vertical creeping
with and without loads 33
G. The effect on locomotion of
de-eying Helix 43
Behavior of the detached foot 49
E» Effect of mechanical stimulation
and of Ringer's solution on
locomotion 51
F» Action of adrenalin on the
neuro-muscular activities
during locomotion 62
Effect of strychnine sulphate
on locomotion 98
CHAPTER VI Summary and conclusions 115
CITATIONS 118
INDEX OF FIGURES
123
1
Chapter I.
Introduction
Precise measurement of vital processes is necessary in
the investigation of many of the problems arising in the
study of vital dynamics. Results based on series of facts
obtained upon the intact living organism prevent confusions
which arise when less exact methods are employed. The
philosophical approach into the realm of vital phenomena
has not been a step forward because the introduction of
philosophical ideas, such as adaptation, behavior, emergence,
psychobiology and purpose have resulted only in methods of
vagueness and inexactness.
Huxley (1854) predicted that the science of the biologist
would be as deductive and exact as mathematics. He was of
the opinion that since biology is a physical science the
methods of this science must be analogous to those followed
in other physical sciences. This prediction was fulfilled
sooner than he would have believed possible. Loeb (1888)
laid the foundations for this method of precise measurement
by his contribution of tropistic conduct. Crozier (1928)
has taken tropisras as a working tool for ttie further devel-
opment of many unexplained activities found in living phen-
omena. He recognizes the significance of the quantitative
aspects of behavior and believes that it is this quantita-
tive treatment of tropistic behavior which is essential if
an understanding of conduct is to be furthered, ^.s he has
I
2
stated, the biological system presented by a sin^^e individ-
ual is not a "thing", a single event, but a system of rela-
tions. These relations must be defined through investigatio!
and their functional dependence analyzed.
Stier (1928) has mentioned certain aspects of animal
conduct other than those considered in the tropism doctrine;
that is, motor activity and fluctuations of sensitivity may
be based on principles originating in the fundamental laws
of chemistry and physics.
Rose (1929) emphasized the fact that it is only by
more precise measurement that the problem of vital dynamics
can be cleared up, and its progress can be furthered only
by the use of quantitative methods and rigorous analyses.
By delicate measurements scientists have obtained just such
desired results, which have completely revolutionized
researches in the field of plant tropisms and at the same
time have opened up new aspects of the physiology of plants.
Furthermore, various related phenomena of animal reac-
tions cover a wide field. Myogenesis (Carey, 1919-20),
oxidations as a function of temperature (Crozier, 1924-25),
tonic immobility (Hoagland, 1927), spontaneous movement
(Stier, 1928) and inheritance (Crozier and Pincus, 1929-30)
are a few activities taken from the field of animal conduct
that have led to important results by the application of
the method of precise measurement. For example, Crozier
and Pincus have arrived at a definition of gene differing
from the usual one; for them it is a definition of the
3
effect in inheritance as a function of some controlling,
independent variable.
In the investigation of the dynamics of histogenesis,
Carey (1919-20) has shown by the method of precise measure-
ments (the number of contractions made in a unit of time,
measurement of intra-vesicular hydro-dynamic pressure and
volume of stimulus) that the formation of muscular tissue
is due to a definite active process, not a passive one,
as the term self-determination denotes. He proved by his
experiments that the intensional stimulus is the necessary
factor in myogenesis. From his dynamic point of view the
muscle types represent differences in the amount of work
that has been done on the undifferentiated mesenchyme by
the differentially growing parts of the embryo during the
active growth period. He demonstrated that it is possible
to transform unstriated muscles into striated by varying
the velocity of application and the intensity of the ten-
sional stimulus to a higher optimum degree. 3y this taethod
of procedure his results enabled him to conclude that the
variable intensity of the optimum tension determined the
muscular type. In other words, the structure of muscular
tissue is determined by the function it performs and the
work it does, but on the other hand structure does not deter-
mine function. He thus reached these conclusions as a
result of his experiments employing analysis through tension
and pressure.
Therefore, just as analysis through temperature
characteristics plays an important part in the snocific
control of vital processes, so also analysis tiiroufih pres-
sure and tension becomes an additional factor which may
lead to identification of reacting living matter. The fol
lowing experiments are approached through this procedure.
5
Chapter II.
Purpose of this investigation
The purpose of this investigation is the detailed
analysis of the factors involved in locomotion, to eain
additional information concerning the creeping mechanism
of Helix. Inasmuch as the movement of the waves found in
the foot of gastropods is inseparable from locomotion it
is necessary to describe the types of waves found in these
animals; and it is also of importance to consider other
forms of periodic waves found in muscular tissue.
Various kinds of periodic waves occurring in muscular
tissue have been investigated to determine, if possible,
the cause of neuro-muscular activity. The waves which are
present in the peristaltic action of the intestine, the
rhythmic pulsation of the cloaca in holothurians , the pedal
waves found in larvae of slug-moths, the waves found in
Thy one briareus , and the peristaltic locomotor waves of the
tent caterpillar may be cited as examples.
The balance of opinion as to the cause of the rhythmic
contractions of the intestine has been in favor of neuro-
genesis, but there is now considerable evidence of tayogenic
origin, offered by Ivlagnus (1905), Gunn and Underhill (1914),
Alvarez and Mahoney (1922), Gowie, Parsons and Lashmet (1929),
Ascanio and ..Ivarez (1929). Alvarez agreed with some of
these investigators as to myogenic origin of peristaltic
action and believed that a gradient of irritability has been
6
found which may be an Important factor in the directlor
normal peristalsis.
The anal pulsating mechanism of holothurians might be
regarded as constituting another unit in the series of inde-
pendent effectors, such as pedicellariae and spines, which
go to make up the echinoderra meuro-muscular equipment.
Crozier (1916) has investigated the physiological character-
istics of cloacal pulsations in Stichopus moebii Semper. The
results of these experiments are in essential agreement with
the data derived from many previous studies in pulsating
structures, such as those of medusae, ctenophores, the
arthropod heart, and the vertebrate heart and intestine,
e» J the rhythm has a temperature coefficient of the order
of magnitude of that for chemical processes and the relation
of pulsation to the salts of sea water is essentially like
that in other well-known pulsating systems.
The pedal waves found in the larvae of the slug-moths
(Gochlidiidae) are similar in many respects to those on the
pedal surface of gastropods. Crozier (1923-24) has found
that the speed of these pedal waves corresponds almost
exactly with the speed of the pedal wave in Chiton tuberculatus
and is therefore in this respect nore comparable to the raol-
luscan foot than to the peristalsis of the body in the earth-
worm or in caterpillars. He has concluded that the peris-
taltic pedal waves of these animals are to be regarded as "a
derivative of the general peristalsis of ordinary caterpillars,
and as in the 'myenteric reflex' of tlie vertebrate intestine
7
it implies reciprocal innervation." (p.. 328)
When the peristaltic locomotor waves of the tent cater-
pillar were studied by Crozier and Stier (1925-26) it was
found that the frequency of the abdominal waves during ver-
tical ascension was controlled by temperature according to
the Arrhenius equation.
Stier (1928) has come to the conclusion that a proprio-
ceptive mechanism in the body wall seems to control the
initiation of the locomotor waves in Thyone.
Lutz (1930) studied the effect of low oxygen tension
on the pulsations of the isolated holothurian cloaca
(isolated strips from the cloaca of S-tichopus moebii Semper
and ring preparations from the cloaca of Gucuinaria frondosa )
because oxygen deficiency has often been associated with
periodicity and augmentation of response in various tissues.
He concluded that a certain degree of oxygen lack results
in increased activity of the tissue.
Although the exact cause of the initiation of the
rhythmical waves (referred to in the preceding paragraphs)
is not known, it can be seen that by means of investigations
of this nature new factors are brought to light that play
an important part in the control of periodic waves.
To investigate rhythmical waves further the gastropods
offer excellent opportunities, and the same principle when
found may apply to other tissues where periodic waves occur.
The action of the waves found in the foot of gastropods has
frequently been compared to the peristaltic waves occurring
in the stomach and intestine. Jordan (1927) found that the
cerebral ganglia of snails have a quantitative influence
on the peristaltic action of the foot (v/ith its pedal r^an-? i
similar to the autonomic innervation of the stomacn. how-
ever, the use of the foot of this animal is of advantage
compared with the use of other tissues such as excised
pieces of intestine. In the study of the latter many more
factors enter, such as regulation of temperature and of
intra-intestinal pressure, and the use of oxygenated Locke's
solution, as well as other factors, thus adding to the com-
plication of investigation.
Helix pomatia (Pig. Al, a and b) and Helix lactea
(Pig. Al, c and d), common Liediterranean species, were used
in these experiments, which extended over a period of three
years. As the animals possess great tenacity of life and
are unaffected by extremes in temperature they are adapted
to experiments continuing over long periods of time.
Another advantage is their precise negative geotropism.
In the experiments of vertical creeping v/ith and without
loads this tropistic quality insures upward creeping.
Still another advantage of using Helix is the tjj)e of
wave it possesses, spreading over the entire foot. There-
fore, the adhesive power of the foot does not present such
a- complication as in the foot that does not form pedal
waves over its whole breadth. Helix is moreover very
suitable for analysis through precise measurements on the
9
intact organism.
Just as investigation through temperature character-
istics has thrown light on the specific control of vital
processes, so analysis through pressure and. tension becomes
an additional factor that may lead to the identification
of reactions in living matter. As the central nervous
interplay governing the mechanism of locomotion and the
exact cause of the waves is unknown in gastropods, the
subsequent experiments v/ere undertaken to investigate
further these waves and other factors involved during
locomotion under varying conditions. The quantitative
measurements of the activities investigated in these animals
have been studied and their functional dependence analyzed.
1
Crozier and Pilz (1923-24) have stated that it should
be possible to utilize these waves for the study of neuro-
muscular physiology in the intact animals, avoiding in this
way effects due to lack of proper circulation and the like
when isolated organs are used.
10
Chapter III.
Review of the literature
The pedal waves found, in the foot of the gastropods
have been described and in some cases classified by Carlson
(1904-05), Biederraan (1905, 1906), Vies (1907), Parker
(1911, 1914), Olmstead (1917-18), van Riynberk (1918-19),
Crozier (1918-19); 1919-20; 1922-23 a and b) , and ten Cate
(1923). Tlie creeping of Limax maximus has been described
by these writers as a result of a rhyth^nic succession of
evenly spaced, progressive wave-like deformations of the
pedal surface.
Vies (1907) has grouped the v/aves that appear on the
foot of mollusks into two general types, i.e., the "direct"
waves that pass from the posterior to the anterior end of
the foot when the animal moves forward, and the "retrograde"
waves v/hich pass over the foot from front to back as the
animal m.oves forward."^ In both types of movement several
subtypes can be distinguished as determined by the lateral
extent of the pedal wave, i.e., "monotaxic", "ditaxic" and
"tetrataxic" subtypes. The direct waves may be m.onotaxic,
ditaxic and tetrataxic types, the retrograde waves either
With the exception of Chiton gastropods always i.iove
forward and never backwards, regardless of the type of
waves passing over the foot. Parker (1914) has found that
Chiton can reverse its locomotion and creep backwards a few
millimeters. This is the first instance of backward loco-
motion to be recorded in gastropods (verified later by
Crozier and Navez).
11
monotaxic or ditaxic. The direct monotaxic waves consist
of a single system of waves traversing the foot as found
in Limax and Helix. Direct ditaxic waves consist of two
systems of waves occupying each one of the lateral halves
of the foot and alternating regularly on the two sides of
the median line which is not affected by these waves.
Parker (1911) has added Tectarius nodulosus and Nerita
tessellata as ditaxic gastropods and has found that in
these animals the waves on the two sides of the foot
usually alternate and are so extensive that never more than
two waves can exist on one side at one time. As a result
of this the foot moves forward in alternate steps, first on
the right and then on the left, the motion resembling that
of a person walking in a sack. Vies (1907) has described
the direct tetrataxic waves, i.e., the foot which is
traversed by four systems of waves is broadly fissured on
the median line and each lateral sole is overrun by two
systems of alternate waves such as are found in the ditaxic
foot. The retrograde waves may be monotaxic or ditaxic.
In considering the theories advanced to explain the locomo-
tion of gastropods only the first group (direct monotaxic)
is considered by the following authors.
The investigation of the activity of these waves has
resulted in various conclusions concerning both the mechan-
ism controlling them and other factors in locomotion.
In Helix the waves are m.onotaxic, but they pass over the
entire breadth of the foot (helicine foot), whereas in Limax
they appear only in the central longitudinal area of the foot.
12
Contraction of the longitudinal muscles, the action of the
dorso-ventral muscles (transverse and oblique rauscles),
the pressure of substances in the body fluids, and other
factors have been advanced to explain the controlling
mechanism.
Simroth (1878) accounted for locomotion by a theory
that he called "extensile muskulatur". He concluded that
the waves cannot be produced by the separate or the combined
contractions of the oblique and transverse muscular strands,
but that the cause of the extension of the foot is the active
extension of the longitudinal musculature. He believed that
the extensile muscle fibers in all or nearly all snails are
the important factors in its m.ovement from place to place.
Jordan (1901) advanced a theory which he later r.iodi-
fied. He did not accept the theory of "extensile muslrulatur"
in accounting for locomotion in the m^^rine gastropod .^tplysia.
He attributed the relaxation or the extension of the longi-
tudinal muscle of the foot to pressure of isolated bodies
of the visceral fluid or blood.
Bohn (1902) attributed the production of the waves to
direct excitation of the m.uscle fibers and to a sort of
progressive "induction". He explained that this "induction"
would be produced every time the m\iscle fibers v/ere placed
on level cylindrical or conical surfaces and the waves
would be distributed in bands or rinfi;s, parallel, narrow
and identical vath each other. He called then organic
waves and postulated the theory that by the study of these
13
and other orp;?:nic waves other facts of biolop-io i^i^I^'ctlon
may be brought to li,(7,ht and may play an important role in
an explanation of the kinetogenetics of evolution and per-
haps even of heredity.
Carlson (1904-05) has described the external mechanics
of locomotion and has differed in some respects from other
authorities. He did not accept Simroth's theory of "ex-
tensile muskulatur" or Jordan's theory of the pressure of
isolated bodies in the visceral fluid. Carlson tested the
longitudinal muscle in the foot of several gastropods and
his results showed that there is no difference between the
physiology of the muscle and that of any other muscle. He
concluded that during ordinary progression the animal
assumes its greatest length and smallest diameter, due to
the contraction of the transverse and the oblique muscles
of the dorsal and lateral parts of the body. The waves
of locomotion in the foot are diminutive representations
of the waves of relaxation and contraction. At the areas
of relaxation the sole of the foot adheres closely to the
ground and between these points the sole is slightly
elevated. Although von Uexkllll believed that the foot is
provided v/ith some mechanical device such as the setae of
the earthworm, Carlson believed that the area of contact
of the foot with the ground in any region serves as a fixed
point through friction, and acting on this the contraction
of the longitudinal muscles of the foot pulls the neighbor-
ing portion of the body forward.
14
Parker (1911) has described most satisfactorily the
external mechanics of locomotion in Helix pomatia. fie
stated that the forv/ard movement takes place in the dark
waves, and quiescence is characteristic of the intermediate
lighter portions of the foot. Each wave is a pulse of for-
ward motion and the rest of the foot is momentarily quies-
cent. He stated, "the area covered by the v/ave is probably
a fourth or fifth of the wnole foot, any moment, there-
fore, three-fourths to four-fifths of the surface of the
foot is stationary and about one-fourth to one-fifth is
moving forward. In other v/ords, the snail stands on the
greater part of its foot while it moves- forward with a much
lesser part." (p. 102). Prom, experiments performed he
believed that each wave on the underside of the foot (of
flelix pomatia ) is a slight concavity. I'Vhen the muscle of
the foot relaxes the portion of that foot that v/as elevated
is returned to its former level and the muscle recovers
its original length and position. This action of the
dorso-ventral muscles takes place from behind forward and
thus a concave wave runs on the surface of the foot from
tail to head. The forv/ard movement of that portion of the
foot which is temporarily lifted from the substrate is
accomplished by the action of the longitudinal muscles.
The contraction of each longitudinal fiber serves to move
the foot forward as the relaxing wave passes over the foot.
It also extends the relaxing posterior fibers. An important
point in this description is that the dorso-ventral :nuscles
15
play an active part in lifting the foot from the substrate
due to the fact that their dorsal ends, being more firmly
set than the ventral ends, serve as relatively fixed points
and therefore the ventral ends move. The action of the
ventral' end lifts the foot locally and overcomes adhesion
in the given region. In this way each point of the foot is
lifted, moved forward and set down again and thus the foot,
and with it the anim.al as a whole, moves forward.
Although Vies and Bathellier (1920) did not arrive at
a conclusion in regard to the action of the pedal waves of
gastropods they deduced that certain numerical laws held
for them.
ten Gate (1923) was of the opinion that the wave of
contraction is caused by the nervous peripheral net-work.
Grozier and Pilz (1923-24) agreed with the description
of the external mechanics of locomotion as given by Parker
(1911). It appears from Grozier 's investigation of the
neuro-muscular activity of the foot that he did not have at
first conclusive evidence of nerve-net transmission. However,
from the experiments (Grozier and Pilz, 1923-24) performed
on the temperature coefficient for pedal activity in Limax
these investigators found that the velocity of a single wave
must have very nearly the same temperature characteristic
which is found also in another case of nerve-net transmission
(Renilla ) . They found that v/hen work was done at a constant
rate the frequency of the pedal waves is influenced by the
temperature according to the Arrhenius equation, with
^ = 10,700 (Qio 11° to 21°G. = 2.1).
16
Magnus (1924) has shown from the "half animal" experi-
ments performed by Jordan and Uexkflll the.t stretchinr-r of
the musculature in snails can react on the condition of the
pedal ffanglia.
Crozier and Federighi (1924-25£) concluded that the
pedal organ of the slug, although under the control of cen-
tral nervous impulses, is essentially an independent effec-
tor. (See further, page49 of this thesis.)
Cole (1925-26) considered the stimulus for geotropic
orientation and locomotion in Helix aspera to be the tension
of the body muscles produced by the downward pull of gravity
The stim.ulus is received by the proprioceptors of these
muscles .
Jordan (1927) explained the rhythmical transmissions
of stimulation in the snail's foot according to von UexkiJlll'
law (p,243). The foot of the snail, with its nerve net, is
not sufficiently autonomous to be able to carry on creeping
without the help of a ganglion. The stimulation which pro-
duces locomotion is conducted in the beginning through the
nerve-net and through the pedal ganglia with the nerves
radiating from them to the periphery, as to the manner in
which the m.ovement itself comes into existence, nothing
exact is known. A true antagonism such as appears in arthro
pods, annelids and vertebrates is lacking in these animals,
and instead the phenomenon of "viscosoid" tonus is present.
Jordan considered "viscosoid" tonus as a static
phenomenon and as an attribute of a colloid system. In his
I
17
distension experiments on the snail's foot he stated that
the characteristics of these muscles can be reco(=i;nized as
those of a fluid colloidal phase. The rausculature of these
animals possesses attributes which in many connections have
the characteristics of protoplasm. For instance, piiago-
cytosis is present in considerable degree in the same animal
in which "viscosoid" tonus plays so important a role in the
outer muscle layer. In amoeboid movements and phagocytosis
the work is connected with the shifting of the parts and
thence with an alteration of the form. IVith regard to
muscle, therefore, a new form of contraction problem arises.
As soon as the musculature is shortened • through stimulation
nothing more is observed of the fluid phenomenon. The con-
tracted muscle is an almost pure elastic body. In tnis con-
nection Freundlich, as referred to by Jordan, has stated
that in dead colloidal stuffs the conversion of the fluid-
viscoid condition into the elastic state occurs. Hypotheti-
cally one may think, in the case of the muscle of a conver-
sion from the sol to the gel condition. In the latter the
form-alteration would no longer be explicable through the
shifting of the component parts bu.t through a particular
reversible elastic form-alteration of them.
ten Gate (1927-28) has stated that the movement of the
appendages of Aplysia limacina is determined directly by
the pedal ganglia and is not Produced by a chain of ganglia.
As real geotropic orientation of the Helix type of
gastropod takes place only during active prop-ression.
18
orientation, because of its close relationship to locomo-
tion, can "be included in this discussion,"^
(jeotropi'c orientation rather than locomotion has been
studied by Grozier and Kavez (1930). They concluded that
orientation is not governed either as to direction or as
to latent period by the involvement of the statocyst.
Gravitationally excited orientation is probably controlled
by the proprioceptive stimulation through impressed tensions
(Grozier and Navez, 1930). As orientation and progression
are so closely united these investigators have stated that
it is still a question as to whether (1) the creeping move-
ment and postural movement are released together or (2)
the operation of the pedal wave is the necessary factor
to act on the stimulus for orientation. Therefore it can
be seen from various theories existing v/ith regard to this
subject that it is important to investigate the question
further. Helix, rather than Limax was chosen, as more work
has be-en done on Limax (Grozier and Pilz, 1923-24; Grozier
and Federighi, 1924-25£) and very few data have been ob-
tained for Helix.
The muscular mechanism for locomotion and that for
orientation is structurally distinct, i.e., creeping is
brought about by means of a pedal organ, wxiile orientation
is accomplished by means of muscles in the dorso-lateral
regions of the body wall.
19
Chapter IV
Method and Procedure
Specimens of Helix lactea and Helix pomatia used in
these experiments were kept at room temperature in separ-
ate jars which were kept slightly moist. Lettuce v/as fed
every other day and the jars were thoroughly cleaned. In
this way the animals lived for six or seven months, the
vigor of creeping showing no diminution at the end of
that period.
The following experiments were carried on within the
temperature lim.its of 19.5° and 24.8°C. The animals were
subjected to varying environmental conditions, which will
be described subsequently. The required measurements were
obtained as follov/s; the animal was placed in a glass jar
1
one and one half feet high and when the snail began to
creep vertically upwards, the sides of the jiar being kept
moist to promote active creeping, the distance travelled
while ten waves were passing over the foot from posterior
to anterior end was measured in millimeters on the outside
of the jar. (Pig. 1). The time was recorded in seconds
or minutes. This constituted one run, and ten runs made
up a series (see Table I). Several series for each animal
were obtained and the conditions for each series were the
^ In later experiments a glass plate set in a frame
similar to a picture frame was used, as this enabled
handling the animal more easily.
20
same. The length of the foot during each run was also
measured in millimeters.
Prom these data it is possible to determine:
(1) The frequency of the pedal waves (F).
(2) The velocity with which a single wave traverses
the foot (v).
(5) The speed of creeping (V).
(4) The advance due to a single wave (A).
An example of the way in which the data observed have
been used to obtain the absolute values of the frequency
1
of waves and the velocity of progression is presented
in detail in Table I, and a description of the way these
absolute values are used in plotting one relation to
another is given in the text.
As the ultimate purpose of this series of experiments
was to analyze the relation between the frequency of waves
and the velocity of progression it was necessary to obtain
from the data thus observed the values of the velocity of
progression, which are given in Table II. With these
measurements it is possible to show by their actual scatter
and by comparative graphs the linear relationsiiips between
the different activities involved in locomotion. The
method of plotting was to take the computed value of the
frequency of waves and the velocity of progression (Table II)
The same method was follov/ed to obtain the measurement
for the advance per wave and the velocity of a single pedal
v/ave.
I
21
and plot the actual scatter, the ordinates representinp;
the frequency of v/aves and the abscissae the velocity of
progression. After presenting the actual scatter in this
way their average points were obtained. The best fitted
line v/as then drawn through these average points. The
graph resulting from the data just given in detail is
shovm in Pig. 3 for animal Ko. 7. Graphs shov/ing the same
relationship have been com.pared in subsequent experiments
when the data are derived under varying conditions.
Tables Ila and lib represent the data for a complete
experiment when each one of the absolute values v/as deter-
mined, i.e., velocity of progression, frequency of waves,
velocity of a single pedal wave and the advance per wave.
From these data the various graphs v/ere plotted as described
above.
22
TABLE I
Observed data"
Calculated
values of
frequency of
waves from
observed data
No. IIo. Ko. Actual Actual Length of Frequency of
of of of time in distance foot during wavcg-^^ Y^^
runs series waves seconds
m mm.
each run^
time
1
10
16.0
7.5
49.0
0.624
2
10
10.8
7.5
51.0
0.925
3
10
13.9
7.0
51.0
0.719
4
10
13.9
8.0
52.0
0.719
5
6
10
= 1
10
12.4
11.2
8.5
7.5
54.0
53.0
0.806
0.892
7
10
12.2
10.0
53.0
0.819
8
10
12.0
7.5
54.0
0.832
9
10
11.4
8.5
53.0
0.878
10
ro
10.4
10.0
54.0
0.961
^ Anj.mal TIo. 7.
2
These measurements are needed v/hen the velocity of a
^ ^ . ^ • tr Length of foot
single wave is to be determined, i.e., V = t^^q for i wave *
23
TABLE II
Calculated
values of
progression
1 obtained from
Observed data observed data
No, No. No. ^LCtual j.ctual Length of Velocity of
of of of time in distance foot during .procuress
runs series waves seconds in mm. each run = di stance ("im_.J,
tin-:e ( sec . )
1
10
16.0
7.0
49.0
0.437
2
10
10.8
7.5
51.0
0.694
3
10.
13.9
7.0
51.0
0.503
4
10
13.9
8.0_
52.0
0.574
5
10
12.4
8.5
54.0
0.685
= 1
6
10
11.2
7.5
52.0
0.669
7
10
12.2
10.0
53.0
0.819
8
10
12.0
7.5
54.0
0.624
9
10
11.4
8.5
53.0
0.745
10
10
10.4
10.0
54.0
0.961
Animal No. 7
24
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26
Chapter V.
Results and Discussion
A. Verti.cal ascension without added load.
In a series of experiments extending over a period
of several months eleven animals. Helix lactea, of vary-
ing v^eights v/ere used. Measurements were obtained for a
total of 36 series, each series made up of ten runs, and
each run made up of ten waves; thus 3600 waves were ob-
served.
Data were obtained from observing 400 waves in each
of the animals when creeping vertically upv/ards carrying
no load. The relations betv/een the velocity of a singl'e
wave and the velocity of progression, and between the
frequency of waves and the velocity of progression, as well
as between the advance per wave and the velocity of pro-
gression were analyzed.
Fig. 2^ shows the relation
a single pedal wave and the rate
It can be seen that a linear relationship exists regardless
of the weight of the animal.
The actual observations were plotted taken from data
obtained from 4 series of 40 runs, each run consisting of
the record of the time and distance covered for 10 waves.
In order to obtain the average points of Fig. 2 the mean
was taken of the actual observations. This was the way in
which the points were determined in the other figures. It
is possible to obtain the average deviation from the mean
for any of the points given in the figures.
between the velocity of
of creeping ^,
i
27
Weight Slope of line"^
Animal Ko. 1 6.5 gm. 0.20
" "4 8.6 " 0.36
" " 7 7.9 " 0.34
Fig. 3 shows the relation between the freauency of
) . It can
waves and the velocity of nrogression (F
V
he seen also that a linear relation exists regardless of
the weight of the animal.
Weight Slope of line
Animal Ko. 1 6.5 gm. 0.20
" " 7 7.9 " 0.18
" "10 3.8 " 0.20
Pig. 4 shov/s the relation between the advance per
To find the slope of the line as given in Fig. a, the
distance AB expressed in mm. is subtracted from the dis-
tance CD expressed in mm. and divided by the distance
represented by line AG. The same unit for AG is used in
each case, i.e. , 50 mm.
28
The results
wave and. the velocity of pro/3;ression (^ )
V
show that in this relationship also a linear rele tion3xj.ip
exists regardless of the weight of the animal.
Weight Slope of line
Anirfial Ko. 4 8.6 gm. 0.5
" " 7 7.9 " 0.3
" " 10 5.8 " 0.3
Further experiments of this nature were performed on
a number of different animsls to see whether consistent
results v/ould be obtained. The following summary shows
the scope of the experiments from which data were obtained
for the three different relationships: V
V V
A for animals 1, 3, 4, 5, 6, 7, 8, 10 and 11
V
during vertical ascensions without load. For these nine
animals a total number of 3200 waves was observed and
measurements taken. From these data graphs were plotted
of the actual scatter for each animal showing the relation
between the velocity of a single pedal wave and the rate
of progression (V j ), betv/een the frequency of waves
V
and the rate of creeping (F
) and betv/een the advance
). The
V
per wave and the velocity of progression (A
V
average points of each graph v/ere obtained and the best
fitted line dravm. Fig. 4A shows a comparative graph of
these lines, each line illustrating for each animal the
relation between the velocity of a single wave and the
29
velocity of progression. A straight line relationship is
found to exist under these conditions. The slope of the
line for each animal was determined. The same method of
comparison and analysis was employed in the investigation
of the relation between frequency of waves and the velocity
of progression (Fig. 4B), and "between the advance per wave
and the velocity of progression (Fig. 4C ) . These results
are given in Table III. ^
Kelix pomatia was used in a series of experiments to
determine whether the laws derived from observations of
Plelix lactea would be confirmed. The conditions and
method of these experiments were the same as for Helix
lactea. The limits of temperature v/ere 16.2 to 21.0°C.,
but constant for each experiment. The length of the foot
of Helix pomatia vmen creeping is from 60 to 100 milli-
meters (Figs. Ala and Alb), and that of Helix lactea from
25 to 40 millimeters (Figs. Ale and Aid). Helix pomatia
has the helicine foot, i.e., it forms waves over the whole
breadth of its foot, so that adherence and progression is
not attended to by functionally separate divisions of the
pedal surfaces. Table Ilia"'' shows some average results
of these experiments. Fig. 4D illustrates that for Helix
pomatia a linear proportionality exists between the velocity
of a single wave and the rate of creeping. Fig. 4E shows
that the frequency of waves is directly proportional to
1
Tim-e is recorded in minutes.
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31
the rate of creeping. Fig. 4P demonstrates that the advance
per wave is directly proportional to the rate of creeping.
Analysis of the experimental results obtained in this
series yields certain information regarding the condition
of the foot during vertical ascensions without load. A
study has been made of the relations betv/een the velocity
of a single pedal wave and the velocity of progression,
between the frequency of waves and the velocity of pro-
gression, and between the advance per wave and the velocity
of progression. The slope of the line for these relation-
ships has been determined. A comparison of these slopes
shows the deviation of the linear proportionality, if any.
Variations from strictest proportion found occasionally
may arise in large part from the complex character of the
pedal musculature. In general the linear proportionality
of these relationships to each other is unmistakable.
The relation between the velocity of progression and the
veloclt^r of a single v/ave shows the greatest alteration
during a given number of runs under the same conditions
for a number of animals of varying weights. The relation
between the frequency of waves to the velocity of progres-
sion exhibits the least alteration within a number of series
of one anima]. or when several animals are compared with
each other, study of the graphs and tables shows that
the slope of the line in a given relation may be the same
regardless of the iveight of t-"'e rrimal.
Therefore, these results Siiov/ b-.at the speed of
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33
creeping for Helix lactea and Ilellx pome-. t la without load
on a vertical surface during upv/ard progression is directly
proportional to the velocity with which a single average
wave courses over the foot. It is independent of the
weight of the animal and the length of tlie foot. ITie
velocity of progression is directly proportional to the
frequence of waves and to the advance of a single wDve.
B. Vertical ascension v/ith added load.
The technique for adding the loads was to attach to
the shell lead v/eights held on by pieces .of adhesive tape.
The animals (Helix lactea ) were first tested to determine
the weight necessary for complete exhaustion. This was
for the purpose of gaining an indication of the probable
number of weights to be attached and to select individual
weights that could be added conveniently. Table IV
represents the results when the animals v/ere tested for
complete exhaustion.
TABLE IV
No. of Animal Complete Exhaustion Weight of Animal
v/lth v/eight
1 4.3 gm. 6.5 gm.
2 0.7 " 7.1 "
3 3.5 " 7.7 "
4 4.8 " 8.6 "
5 3.0 " 7.1 "
6 4.8 " 6.7 "
7 4.8 " 7.9 "
8 3.8 " 8.5 "
9 2.0 " 3.5 "
10 4.8 " 3.8 "
11 3.0 " 5.0 "
I
34
Animals Ko. 2 and Ko. 9 showed the most rapid exhaustion
with the least load. Some inherent weakness may have
accounted for this as both animals died after three months.
Animal l\o, 2 crept poorly from the beginning and it was
possible' to obtain onl^;- half the number of runs as com-
pared with the other animals when creeping v/ithout loads.
The same v/as true for No. 9, except that great activity
was shown for the series that was obtained, though only
half the series as compared with the other animals was
obtained. The results show that exhaustion caused by
added weights was independent of the v/eight of the animal.
The load which caused exhaustion for the majority of the
animals v/as 3.0 gm. or more up to 4.8 gm.. Conseauently
individual lead v/eights of 0.5 gm. v/ere chosen as the
most suitable to obtain effects of small added loads up
to 4.3 or 4.8 gm.
The method of applying the weights was as follov/s:
the weight for each load v/as gradually increased but these
increased loads were not added in immediate succession.
A run without any added load or a load smaller than the
load just used was made between the various increases.
Table V illustrates this method. The advantage of this
method was to insure the fact that any effects of fatigue
caused by the additional weights could be avoided by having
a run of no load or a lighter load than had just been
removed before the next heavier weight had been added.
This was the method used to obtain data for eleven
animals, the ranpie of temperature for the entire series
being from 19,5° to 24.8°G. but constant for each e:x:peri .lent .
The way in which these graphs were made was to take
the data from a group of animals, i.e., animals numbered
3, 4, 7 and 10 and use it to plot four different comparative
graphs (each graph for each animal) showing the following
relations :
a. Between velocity of progression and velocity of
pedal wave.
b. Between freouency of waves and velocity of
progression.
c. Between advance due to a single' wave and speed
of creeping.
The measurements obtained vmen 8,300 waves were ob-
served while eleven animals were carrying varying loads
during vertical ascension were used in the analysis of the
data. A comparison was made with and without load from
the actual scatter of the data. The same relationships as
those described for the last experiments were used as
functions of one another. Collective graphs were plotted.
The following typical graphs illustrate the way the data
v/ere utilized throughout these experiments. Fig. 5a shows
the actual scatter obtained from direct observations from
a series of runs with and without load for animal l\o. 7.
The relation between the velocity of progression and the
frequency of waves has been plotted. Fig. 5 also shows
the effect of added loads on animal Ko. 7 'vhen the freouency
I
56
TABLE V
Method of obtaining data v/iien varying loads v/ere added
to one animal during vertical creeping.
Animal Series 10 Runs of Without Kdded 'jVith Added
10 Waves each Load Load
1 "
tt
2 "
0.7
gm.
(run
1)
w
4 "
1.2
tt
(run
1)
5 "
0.7
11
( run
2)
6 "
2.0
ji
7 "
ti
8 "
2.5
II
9 "
0.7
II
(run
5)
10 "
5.0
tt
11 "
\»2
ti
( run
2)
12 "
5.8
II
15 "
14 "
4.5
tj
etc. until
exliaustion
37
of waves is plotted against the velocity of progression.
However, this graph has been derived by first plotting
the actual scatter for each added load (each load plotted
on a separate graph). Then the average points of the
actual scatter for each load were determined and the best
fitted line drawn. A comparative graph v/as made of these
lines - each line representing an added load. Fig. 53
also illustrates for animal No. 10 the relation between
frequency of v/aves and the velocity of progression when
creeping vertically upwards carrying varying loads.
From an analysis of the data obtained under the
above conditions it is possible to study the individual
effect of each load on the various relations. This indi-
vidual effect can be observed by comparing the slope of
each line for each load. For instance, in Fig. 5 these
results show that this relationship (frequency plotted
against velocity of progression) is independent of the
load carried. This is strikingly illustrated when we
analyze these slopes; for exam.ple Mo. 7, without load has
a slope of 0.20, with 3.8 gm. added the slope became 0.22,
and with 4.3 gm. it was again 0.20.
Pig. 6 illustrates the effect of tension as related
to velocity of progression. No definite lav/s can be
stated, but in general the first loads (0.7 gm. to 2.0 gm. )
tend to increase the velocity of progression. A load of
2.5 gm. tends to diminish the rate of creeping and v-ith
increasing load usually there is a sli^uit increase in the
58
rate and then no change.
After the results were ueterti\lr.t.a as p;iven in Fip;. 6
a comparison was made to determine the probable error
when plotted with var^z-ing loads. Factors v/hich account
for error may be fluctuations in the velocity with v/hich
the individual waves traverse the foot and the possibility
that the adhesive pov.er of. the foot largely depends on its
extruded mucous material v/hich may result in a slight
error when comparing the effectiveness of the pedal mechan-
ism.
The probable error of any one of the activities may
be plotted against varying loads and then ' compared v/ith
the normal. graph showing the relationship between the
probable error for velocity of progression and varying
loads is given in Fig. 7"^. The probable error appears to
be greatest when weights of 0.7 gm.. , 1.2 gm. and 2.0 gm..
were added, and very slight with 3.0 gm. , 3.8 gm, and
4.3 gm. A large probable error in experiments of this
kind would not be expected because the factors observed
in these experiments are very definite, each wave being
readily discernible.
Using Bess el's formula for probable error;
0.6745 , ~v
/\Jn (n-1)
the- follov/ing values were obtained and plotted against the
varying loads. Animal Ko. 6.
V/ithout loads 3,04 With load 2,5 gm. 5.12
With load 0,7 gm. 4,34 , " " 3.0 " . 3.45
" " 1.2 " 3.96 " " 5.8 " 5.35
'» " 2.0 " 3.50 " " 4,5 " 5.45
39
Table VI shows the effect of loads on the individual
factors involved in locomotion, i^bsolute values and' per-
centage of increase or decrease during vertical ascensions
with added loads are compared with ascensions v/ithout
load. In analyzing the effects of these added loaas on
the individual activities the freauency is found to snow
the least alteration and velocity of progression the
greatest.
The results of the foregoing experiments confirm for
the first time for Helix these facts relative to loco;7iotion.
Tiie results show that the velocity of a single wave, the
frequency of the waves and the advance per wave as related
to the rate of creeping are independent of the load carri3d.
These facts show that the foot is an independent effector*
This evidence indicates that the intrinsic neuro-muscular
mechanism of the foot is the primary factor in locomotion.
Cole, on the contrary, has stated (1925-26) that the
stimulus for locomotion in Helix is tension of the body
muscles produced by downward pull of gravity and that the
stimulus is received by proprioceptors of these ::iuscles.
This conclusion would indicate that the primarv fpctor in
locomotion is the central nervous control v/hicu coes not
accord with the experimental findings of the above experi-
m.ents. Experiments reported later in this thesis show
that the foot is secondarily under the control of central
nervous" impulses but that primarily it is an independent
effector.
40
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45
C. Vertical ascensions of the de-eyed Helix with and
v/ithout loo.d.
A series of experiments was carried on to determine
whether the lav/s of linear proportionality governing loco-
motion would be altered if the animal were de-eyed, hs
far as can be determined no mention has been made in the
literature of experiments of this nature. A comparison
was then made between the normal animals and the s&rn.e ani-
mals de-eyed (Helix lactea ) . The eyes are easily renoved
as they are situated at the tip of the posterior tentacles
which are supplied by nerves from the supraoesophageal
ganglia, i.n alteration in the velocity of the pedal waves,
the velocity of .progression, the frequency of waves and
the advance per wave, even though slight, might be expected
in the de-eyed animal v/hen the posterior tentacles are
cut off. De-eyeing should interfere with reflex interplay
due to the connection of the nerve net found in Helix.
iiervous system in Helix. In the nervous system of mol-
luscs there are some highly characteristic features. In
Helix it consists of three pairs of ganglia associated with
important sense organs and connected by nerve cords. One
pair (Fig. A3,c) v/hich lies dorsal to the oesophagus supplies
the tentacles and the eyes. Sometimes this cerebral ganglion
is considered to be made up of two supraoesophageal ganglia
joined by a broad transverse commisure. A second pair (Fig.
A3,p) lies TFciitral - to the alimentary tract on the front part
of the muscle mass of the foot. These are the pedal ganglia
which are connected with the otocysts, consisting of two
small sacs imbedded in the pedal ganglia. The third pair,
the visceral ganglia (Fig. A3,v) are also ventral. The
arrangement of the nerve-net can be seen from Fig. A2.
44
The supraoesophageal ganglia are connected with the sub-
oesophageal ganglia, which consist of tv/o principal
ganglionic masses. The forward mass is a pair of ganglia,
the pedal ganglia, and the posterior mass consists of the
visceral ganglia. Therefore, since the supraoesophageal
and suboesophageal ganglia are connected it is expected
that de-eying v/ould affect the reflex control of the pedal
ganglia.
After measurements were obtained for the normal animals
with and without loads v/hile creeping vertically upv/ard,
the same animals were de-eyed. The temperature varied
from 19.5*^ to 24.8°G. but v/as constant during each experi-
ment."'" Data were gathered from the observation of 1100
waves for the normal and the de-eyed animals carryine- no
load. For animals carrying various loads 2700 waves «ere
obtained and measurements taken. Graphs showing the
actual scatter obtained from these measurements and other
graphs showing the average points of the actual scatter
were plotted according to the m.ethod described for the
previous experiments.
Table VII shows a comparison of the de-eyed and normal
anim.al during vertical ascension with and without loads.
Absolute values and the percentage increase or decrease
are given for the velocity of a single pedal wave, the rate
of creeping, the frequency of waves and the advance per
wave. In a great many cases the individual activities
V.Tierever temperature limits are m.entioned it is to be
understood unless otherv/ise stated that the temperature was
constant for each experiment.
45
taking place during locomotion in the de-eyed animal are
increased when compared with the normal. This increase
may be caused by the removal of the eyes. Reference to
Pigs. 8, 9 and 10 shows that the linear relation betv/een
velocity of progression and frequency of pedal waves or
that between speed and dimensions of a sing-le wave, or
between the advance per wave and the rate of creeping is
not altered in the de-eyed Helix lactea .
These facts are further illustrated by Figs* 8A, 9A
and lOA. These shov/ the actual scatter obtained from direct
observation of the normal and de-eyed animal. The relations
between the velocity of a single pedal wave and the rate
of creeping (Fig. BA), betv/een the velocity of progression
and the frequency of waves (Pig. 9A) and between the
advance per wave and the velocity of progression (Pig. lOA)
have been plotted for animal No. 3 during vertical ascen-
sions v^ithout load.
It was determined also froro a series of experiments
that the carrying of added loads during vertical ascen-
sions does not alter the linear proportionality between
these relations in the normal and de-eyed animal. Figs.
lOB, lOG and lOD show the actual scatter obtained from
direct observations under these conditions (normal and
de-eyed). The relation between the velocity of the pedal
waves and the rate of creeping (Fig. lOB), between the
velocity of progression and the frequency of waves (Fig.
IOC) and between the advance per v/ave and the velocit^^ of
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46
..oeression (Fig. loD) has been plotted for animal No. 3
enuring vertical ascensions with added loads of 0.7 grama.
Fig. lOE and Table Vila illustrate that the linear
relation between the velocity of a single pedal wave and
the rate of creeping is not altered when the de-eyed
animal (ITo. 4) is creepin.. upward carrying added loads.
TABLE Vila
Animal No. 4 Added load Series slope of line
0.7 gni. 1
0.7 " 2
0.7 " 3
1.2 " 1
0.26
0.24
0.24
0.52
0.24
1.2 " 2
^•^ " 0.26
Q C ft
0.30
^•^ " 0.24
3«8 " 0.14
" 0.64
exhaustion
These data show that the foot is primarily an
Independent effector. It may be secondarily under cen-
tral nervous control as evidenced by the slight alteration
49
that occurs in the Individual activities wnen the eyes are
removed.
The significance of the terra "independent effector"
is shown by the fact that "for equal rates of creeping
(Liiaax) the activity of the pedal organ is independent of
the added loads. This is consistent .vith the fact that the
activities Of the pedal organ are determined by its intrinsic
neuro-muscular structure. The nervous elements (nerve-net)
in the foot are secondarily under the control of the central
ganglia." (^rozier and Federighi, 1924-85,0). Added loads
do not alter the intrinsic activity of the creeping organ,
but appear to act on the central reflex mechanism which
inhibits or releases the pedal waves. Therefore we may
speak of the foot so far as it concerns its production of
pedal waves as an "independent effector" secondarily under
the central nervous control, since the laws of its aotlvit-
are the same regardless of the load carried in locomotion.
D. Investigation of the beliavior of the detached foot.
The foot v/as detached from the pedal ganglia to gain
further information concerning the ad jus tor mechanism
regulating locomotion. diagram showing (a) the nerve-
net in the foot of Helix pomatia and (b) the principal
parts of the nervous system is snown in Fig. 2.., taken from
Ubujigen aus der vergleichenden Pliysiologie", by Jordan
(1927, p,22.5). By comparing the normal v/ith the pedal-free
50
anl^l Jordan h.s concluded that the principal lur.ctlon of
the pedal gan^c^ia as v,ell as the sin^cle ganglion of the
.-acidlana is the reg,.lation of "vlsoosoid" tonus, i-e con-
cluded that the uniform condition of the musculature in
the nornial animal is to be considered a sort of an eauill-
brium between two processes, the peripheral and presu,^,bly
reflex Production of resistance, and the steady lessening
of this condition by the pedal ganglia.
The following experiments on twenty-four animals were
performed to study tne oenavior of the foot when separated
from the pedal ganglia. A series of anim.ls was used in
Which the foot n.d neen cut directly fror. .ne visceral
organs. Immediately after the snail had been creeping
actively the foot was held aw,y from th. shell by means
of a forceps placed as near t.., visceral organs as possible,
and then the foot was cut off quickly.
The isolated foot -.vas cttacied at one end by silk
thread to c. ..-ar;; lever ana at the other end to an L-snaped
rod (Fig. 11) which was held in place by a muscle-clamp.
A moist chamber surrounded the foot. J.ymograph records
were ootained snowing the action of the foot. Tne limits
of temperature for the entire series of experiments were
from IS. 5° to 24.8°C;.
Reference to Figures 11a &nd lib shows that the usual
rhythmical action of the foot is rre^.tl-r interfere-
when it is not conneote. .vlsn tae pedal ganglia. Even in
kyinograms showing some recurring waves the usual periodic
51
recurrence found In the intact animal is destroyed. The
destruction of the periodic recurrence of .avea when tne
pedal ganglia are separated from the foot suggests that
this periodicity is determined by the presence of these
ganglia.
Extirpation of the pedal ganglia in the intact anirnal
may be possible and further study of the locomotor waves
under these conditions may bring to light additional infor-
mation as to the exact action of the pedal ganglia in this
neuro-rnuscular activity.
E. The effect of mechanical stimulation and of Ringer's
solution on locomotion.
Preparatory to experiments Involving the injection of
certain drugs a series of control experiraents ..vas performed
on Helix lactea and Helix pomatia. The controls ,vere tested
for (a) consistency of runs, (b) the effect of mechanical
stimulation, (o) the effect of Klnger's solution (cold
blooded) .
In testing the consistency of the runs a comparison
was made of the data obtained for runs ^ separated by intervals
of time. Reference to Table VIII sho^.vs that in a ..riven ex-
periment the data obtained at the end of three hours may
vary slightly from the data obtained at the end of one or
^ The composition of Ringer's solution (1000 cc. of --ter)
used m every case was as follows- .^ter)
Grams
i:aCl 6.5
KCl 0.14
CaClg 0.12
I^'aHC03 0.20
52
at the end of two nours. Except fop this slir^^t variation
the results ms.y be said to be consistent.
The controls ivere also tested to determine the mechan-
ical stimulation on vertical creeping. mien tlie animal
was creeping actively upwards a hypoderr.iic needle, v/hich
was to act as the mechanical stimulus, was inserted into
either the body v/all in the region where the visceral mass
joins the foot or into the anterior end of the foot ^.ir-ctlv
back of the tentacles. The reactions of the animal were
noted.
The method followed in these experiments is illus-
trated by the follov/ing, v.-hich represent the procedure of
a typical experiment in this series v/hen the animal was
creeping upward.
(a) Records of norm.al run obtained.
(b) Records taken of runs after mechanical stimulation
of the body wall.
(c) Interval allowed to remove the effects of this
stimulation.
(d) Further records of normal run.
(e) Records of runs obtained after mechanical stimu-
lation of anterior end of the foot.
In eight out of ten animals used the mechanical sti:au-
lation of the body wall caused a withdrawal of the head
into the shell, followed by a slight contractio:-^ -^f -^':e
foot; then creeping began immediately. In one of the other
two cases (experiment 50, animal No. 15) creeping did not
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55
begin for 43 minutes after stimulation. In the other
instance (experiment 52, animal IIo. 16) it did not begin
for nine minutes after pricking.
Llechanical stimulation of the body wall also caused
an increase in the velocity of progression and in the ad-
vance per wave in Helix pomatia (Table Villa), ar^ either
a decrease or an increase in the frequency of waves and
in the velocity of a single wave. No definite conclusion
can be drawn, however, from this type of experiment on
Ilelix lactea (Table Vlllb), for in some of the experiments
these activities were increased, while in other experiments
they were decreased.
Table VIIIc shows the effect of mechanical stimulation
at the anterior end of the foot does not differ greatly
from, stimulation of the body wall. Here again the results
do not show any definite consistency, the frequency of
waves and the velocity of a single wave being least affecte
Therefore the mechanical effect can be eliminated in the
subsequent experiments.
The effect of Ringer's solution was tested on the
controls. Experiments were carried out by injecting 0.2
cc. of Ringer's solution and studying the effects. Data
for these experiments are given in Table Vllle.
As the chemical composition of the blood of inverte-
brates approaches that which we find in the lower verte-
brates, amphibian (or cold-blooded) Ringer's sol.;;.ijn c_us
the drug was used in the following experiments.
56
TABLE Villa
The effect of mechanical stimulation of the body wall
of Helix pomatia. Absolute values are given for the fac-
tors involved in locomotion.
Ko. of Velocity of
i-'requency
Velocity of
iidvance
Experiment progression
oi waves
single v/ave
per wave
in mm. per
per m.inute
in mm. per
in mm.
minute
iriinut e
5 (Normal)
8.5
42.0
3353
0.24
6 (Mechanical
stimulation)
14.8
43.0
3525
0.33
10 (Normal)
19.2
71.4
6900
0.26
11 (I.iechanical
stimulation )
23.0
62.0
6630
0.56
14 (Norm.al)
13.4
59.5
82i00
0.22
15 (Mechanical
stimulation )
19.0
52.0
7300
0.36
20 (Normal)
12.2
62.5
7600
0.19
21 (Mechanical
stimulation)
18.6
83.3
6700
0.22
57
TABLE VII lb
of
The effect
Helix lactea
of mechanical stimulation on the
, on vertical creeping.
body wall
No. of Velocity of
experiment progression
in mm. per
minute
Frequency of
waves per
minute
Velocity of
sin^f^le wave
in mm. per
minute
i-.dvance
per t»ave
in mra.
41
42
(Normal )
(Lie Chan ical
stimulation )
22,0
27.7
62.5
62.5
4500
4300
0.35
0.44
48
\ IV X ilicL J- 1
(i-iechanical
stimulation)
24.0
62.5
83. 0
4900
4700
0.24
0.29
49
50
(Normal )
(Mechanical
stimulation)
23.0
15.0
71.0
83.0
5700
5100
0.32
0. 19
51
52
(Normal)
(Mechanical
stimulation )
48.0
40.0
83.0
71.0
4900
5100
0.57
72
75
(Normal)
(Mechanical
stimulation)
20.5
15.3
83.0
83.0
4400
4000
0.24
0.18
78
79
(fiormal )
(Mechanical
18.6
r;3.0
4600
0.22
stimulation) 19.6 83.0 5000 0.23
58
TABLE VI lie
The effect of nechanical stimulation of the anterior
end
of the foot
of helix
lactea on
vertical creeping.
No.
of Velocity of
Frequency
Velocity of
Kdvance
experiment progression
of waves
single wave
per wave
in mm. per
per minute
in mm. per
in mm.
minute
minute
43
(Normal )
29.0
100.0
4100
0 . 29
A A
44
(Mechanical
stimulation)
17.0
(DO . \J
40UU
U . cx
47
(Normal )
18.3
83.0
4/00
0 . 22
4c5
(i.Iechanical
stimulation )
l«i) . 8
oo . u
ATy r\r\
4 / UU
0 . lb
53
(Normal )
14.8
71.0
4yuu
U • (dU
54
(Mechanical
stimulation )
22.0
DO . u
U • /i o
55
(Normal )
38.0
125.0
6025
0.30
56
(I.Iechanical
stimulation )
13.6
72.0
4200
0.18
74
(Normal )
15.5
131.0
5550
0.18
75
(Mechanical
stim.ulation )
13.6
72.0
4200
0.18
80
(Normal )
44.0
122.0
6000
0.34
81
(laechanical
stimulation )
20.9
125.0
6375
0.17
TABLE VI I Id
Effect of doses of Rinrer's solution (0.2 cc.
on the velocity of progression (Y), frequency of
waves (F), velocity of pedal waves (v) and the
advance per wave (A).
Kelix lac tea
Plus"^ Minus Cases unchanged
V 9 10 1
P 8 ■ 12
V 8 12
A 6 10 4
Increase or decrease from the normal animal
after injection of Ringer's solution, giving total
number of cases of increase or decrease respectively
60
TABLE Vllle
The effect of Ringer's solution on vertical creeping.
Ko. of
Animal
Velocity of
progression
in mm, per
minute
Frequency of
waves per
minute
Velocity of
single wave
in rnm. per
minute
r.dvance
per wave
in mm.
1
Norma 1
Ringer ' s
38.8
47.4
38.8
37.3
4800
4700
0.36
0.49
2
Normal
Ringer ' s
33,1
38.9
85.4
70.3
5040
5100
0.47
0.52
3
Normal
Ringer ' s
30.9
24.3
75.1
68.6
4450
4370
0.41
0.29
4
Normal
Ringer ' s
20.4
13,1
69.8
65.2
4900
5300
1.09
1.0
5
Normal
Ringer ' s
43.1
25,1
99.4
79.1
5680
5920
0.38
0.31
6
Normal
Ringer ' s
21.1
23.2
68.8
63.8
5550
4440
0.31
0.24
7
Norm.al
Ringer ' s
35.9
22.6
83.5
81.6
5950
5720
0.39
0.52
8
Normal
Ringer ' s
21.0
15.4
70.6
72.1
4200
3880
C.32
0.54
9
Normal
Ringer * s
36.6
39.8
80.1
106.6
5220
4940
0.46
0.37
10
Normal
Ringer ' s
29.4
43.7
58.3
71.1
3980
4720
0.16
0.35
61
TABLE VI He
(cont, )
"Ko . of
Animal
progression
in mm. per
m.inute
rx'ecu.ency
of waves
per minute
veiociLy 01
single wave
in mm. per
minute
i-.dvance
per wave
in mm.
11
Normal
Ringer ' s
9.7
33.7
82.1
57.6
6560
4020
0.39
0.22
12
Normal
Ringer ' s
18.1
29,2
75.0
84.2
4200
4420
0.27
0.35
13
Normal
Ringer ' s
53.8
33.1
94.1
75,9
5140
4520
0.57
0.44
14
Normal
Ringer ' s
27.3
34.1
90,2
91.6
5640
4360
0.32
0,37
lb
Normal
Ringer ' s
28.1
19.3
65,0
66,4
4800
4580
0.28
0.19
± D
Normal
Ringer ' s
19.1
16.1
77,0
61.0
4000
5060
0.21
0.20
JL f
Normal
Ringer ' s
12.7
12,8
69.4
73.3
4800
4450
0.18
0.18
lo
Normal
Ringer ' s
41.1
16.4
82.8
71.5
4560
4900
0.23
0.24
19
Normal
Ringer ' s
29.9
31,1
123.9
97.9
4980
3470
0.48
0.32
20
Normal
Ringer ' s
21.4
18.6
84.7
91.7
4300
4420
0.20
0.21
62
F, The effect of adrenalin on the neui-o-.-nuscuiar activitie
during locomotion.
The action of adrenalin on various kinds of tissue,
particularly on the hearts of vertebrates, has been ex-
tensively studied. Clark (1927) has cited references ■
(p. 62) concerning the effects of adrenalin on some inver-
tebrate tissues, i.e., the intestine of the crayfish, the
oesophagus of i^phrodite, of i-.physia and of Helix. He has
also drawn attention to the fact that the inhibitory
action of adrenalin is as common as its augmentor action
in vertebrates, but that only augmenter action has been
described in invertebrates. Boyer (1926) found that
adrenalin and spartein slackened the rhythm of the isolated
heart of the snails. He found that adrenalin affects
neither the amplitude nor the tonus in the doses used
(1 part in 10,000 and 1 part in 1,000), whereas spartein
reinforces the amplitude and augments the tonus. V/yman
and Lutz (1930) have described an inhibitory •:.c':ion of
adrenalin on the isolated holotnurian cloaca. Their
experiments demonstrated that adrenalin has both inhibitory
and excitatory effects on invertebrate tissues.
Ho reference in the literature as far as can be
determined has been made to any work on the effects of
adrenalin on the muscles involved in locomotion of molluscs
The following experiments describe the action of different
amounts of adrenalin in the intact animal (Helix). Con-
trasted with the study of the detacuea foot, whic:^ i./.o^vea.
63
interference with the circulation, the subsequent experi-
ments show the effect on locomotion, i^drenalin, which
has a low molecular weight, is easily diffusible, is not
attached to protein and is readily poured into the blood
stream from which it is removed by the tissues. Usually
there is no delay in the manifestation of its effect. Its
maximal duration of effect in mam-mals is generally only a
few minutes and its physiological activity during this
interval is narked (Kendall, 1929). However, in cold-
blooded animals the effect of adrenalin m.ay be prolonged.
Bieter and Scott (1929) reported a rise of blood pressure
in Rana pipiens following a dose of 0.2 cc. of epinephrine
hydrochloride, 1:10,000, which persisted for at least one
and one-ouarter hours after injection. MacKay (1931) ■':5.s
found that adrenalin causes rise of ventral aortic jlood
pressure in skates (Kaia) , which persists from one to two
and one-half hours. vVyman and Lutz (1932) reported that
intravenous injection of adrenalin in Squalus acant.-ias
produced a rise of both the ventral and the dorsal aortic
blood pressure, both systolic and diastolic, persisting
for at least thirty minutes. .-.Ithough not a great deal of
work has been done to compare the lasting effect of adren-
alin in the cold-blooded animals v.'ith the effect in m.aminals
the evidence so far obtained shows that there is more pro-
longed maximum effect in cold-blooded animals.
Adrenalin chloride solution (Parke, Davis viV Co.) .vas
used in some of these experiments. In others dry suprarena
64
extract (Parke, Davis a Co.) usea Ir^ oi'aer to check
the results obtained from adrenalin c hloride because of
HGl and chlorotone.
The same general procedure was followed as previously-
described. An investigation was made to see whether the
linear proportionality between the various relationships
for vertical creeping held when adrenalin was injected.
Data were obtained from 68 animals. The results found
for Plelix lactea confirmed those obtained for Helix
pomatia. Therefore, since Helix lactea exhibits more
active creeping and lives in the laboratory for longer
periods of time than Helix pomatia , Helix lactea was
chosen for the majority of experiments.
Data were first obtained for vertical ascensions
under norm.al conditions. Then these data v/ere compared
with the data obtained after adrenalin (of a particular
concentration) had been injected. The data p-iven below
were obtained from a series of eighty experiments. From
the 68 animals used a total of 40,500 waves were observed,
i.e. , 19,700 for the normal animal and 20,800 for the
injected animals. Adrenalin chloride solution (Parke,
Davis & Co.) was added to Ringer's solution to make
various concentrations, i.e., 1:10,000, 1:20,000, 1:40,000,
1:100,000 ,
1:80,000/1:120,000 and 1:200,000. Measurements were first
obtained for vertical creeping under normal conditions.
Then 0.3 cc. of adrenalin (of the concentration to be
studied) was injected into Helix pomatia , and when lielix
65
lactea was used 0.22 cc. of adrenalin was injected. The
time elapsing between the injection and the resumption of
creeping varied in different animals from immediate creep-
ing to one hour. In a few cases creeping did not begin
for two to three hours.
In order to determine whether the linear propor-
tionality between the various relationships for vertical
creeping held when adrenalin was injected the measurements
obtained for the normal and injected animal were employed
as described in the previous experiments. Graphs v/ere
plotted of the average points showing the relation betv/een
the velocity of a single pedal wave and the velocity of
progression, between the frequency of waves and tlie velocity
of progression, and between the advance per wave and the
velocity of progression. The best fitted line was dra';m
and the slope of the line determined. The slope of the
line obtained for a particular relation for the control
was com.pared with the sam.e relation for the injected animal.
Figures 12, 13 and 14 show the typical results. It is
noted that the straight line proportionality and the slope
of the line are essentially unchanged by adrenalin. These
are added features which support the idea of an "independent
effector in muscle.
A description is given below of the effect of
injection of various concentrations of adrenalin chloride.
Table IX shows the effect of 0.2 cc. of adrenalin (1:10,000)
on Kelix lactea during vertical ascensions. i:drenalin of
56
this concentration usually causes a decrease in f^e velocity
of progression, in the frequency of waves, in the vcj-ocI:/
of a single wave and in the advance per wave.
Similar experiments were performed on Helix
pomatia (Table IXa ) . Prom 1950 waves observed on the
normal animal and 2000 on the injected animal data were
procured from, which Table IXa has been computed. Adrenalir.
(0.3 cc. of adrenalin, 1:10,000) had the same effect on
these animals as on Helix lac tea , i.e. , a decrease in each
of the factors involved in locomotion, during vertical
ascensions. Table X shov/s that the same results in the
main are obtained with both species.
Tables Xa through Xh give the data from which
Table X was compiled. Other experiments employing adrenalin
in doses of 1:20,000, 1:40,000, and 1:80,000 (Table XI)
give the same results as obtained for doses of 1:10,000,
i.e., usually & decrease in the velocity of progression,
the frequency of waves, the velocity of a single wave
and the advance per wave. With doses of adrenalin 1:100,000
(Table XIa) inconsistent results were obtained, sonetimes
an increase in the activity and som.etimes a decrease in
the same activity.
However, doses of adrenalin 1:120,000 (Table XI)
produced a stimulating- effect on the velocity of progression,
the frequency of v/aves, ti.e velocity of a si;.;jj.c! .vave •j.U'J.
sometimes either an increase or decrease in the advance
per wave.
67
TABLE IX
Effsct of injection of adrenalin chloride (0.2 cc.
1:10,000) on the velocity of a single wave (v), the rate of
creeping (V), the frequency of waves (F), and the advance
per wave (A) (Helix lactea ) during vertical ascensions. *
Ko. of
animal
linn «
rum.
V =
min.
p _ waves
mm.
— mm.
waves
2
0 = control
• = injected
2576
2682
36,6
34.0
66,0
67.8
0.33
0.29
3
0 = control
0 = injected
3186
3060
45.2
44.7
76.0
67.2
0.38
0.34
7
0 = control
• = injected
5381
4200
30.4
28.1
83.4
80.1
0.34
0.36
9
0 = control
0 = injected
.3334
4704
99.6
55.2
79.2
77.4
0.46
0.36
10
0 = control
• = injected
4660
4358
30.3
23.7
64.6
88.2
0.46
0.29
13
0 = control
• = injected
6080
5250
24.4
25.1
74.3
60.7
0.33
0.42
14
o = control
• = injected
3908
3510
55.3
43.2
79.2
72.0
0.40
0.34
68
TABLE IXa
A comparison of the normal and injected anim.als''' (Helix
poma tia ) during vertical ascensions. Absolute values for
the velocity of a single wave, the rate of creeping, the
frequency of waves and the advance per wave.
Ko. of
animal
Ve
a
in
locity of
single v/ave
mm. per
minute
Rate of
creeping
mm. per
minute
Frequency
in waves per
of
mm.
iidvance
per wave
in mm.
60
Control
Injected
6920
5400
48.2
17.2
96.0
92.0
0.96
0.24
62
Control
Injected
7830
5640
46.8
22.5
103.1
81.3
0.45
0.28
66
Control
In j ected
6010
4950
19.2
36.0
95.2
92.0
0.19
0.38
69
Control
Injected
5770
8800
45.1
29.9
60.0
87.0
0.78
0.50
70
Control
Injected
6280
6460
21.9
19.6
62.2
94.0
0.34
0.31
75
Control
Injected
6350
5210
33.5
18.9
71.9
62.0
0.45
0.32
Injection
of 0.3 cc.
adrenalin
chloride, 1:10,000
•
69.
TABLE X
Effect of doses of adrenalin (1:10,000) on the
velocity of progression (V), frequency of waves (F),
the
velocity
of pedal waves
(v), and the
advance
wave
(A).
Helix
lactea
Helix
pomatia
Plus^
Minus
Plus
Minus
V
1
6
v'
1
5
P
2
5
F
2
4
V
2
5
V
2
4
A
2
5
A
1
5
Increase or decrease from, the normal animal after
injection of adrenalin (1:10,000), giving total number
of cases of increase or decrease respectively.
70.
TABLE Xa
Effect of doses of adrenalin chloride (1:10,000)
on the velocity of progression (flelix lactsa).
l\o. of
Velocity of
Velocity of progression
in mm. per minute
Control
After dose of adrenalin
1: 10,000
2
36.6
34.0
3
45.2
44.7
7
30.4
28.1
9
99.6
55.2
10
30.3
23.7
13
24,4
25.7
14
55.3
■ 43.2
TABLE Xb
Effect
of doses of adrenalin chloride (1:10,000)
on the velocity of progression (Helix pomatia).
llo. of
Velocity of
Velocity of progression
experiment
progression
in mm. per minute
in mm. per minute
after dose of adrenalin
Control
1: 10,000
60
42.2
17.2
62
46.8
25.5
66
19.2
36.0
69
45.1
25.9
70
21.9
19.6
75
33.5
18.9
i
1
71.
TABLE Xc
Effect of doses of adrenalin chloride (1:10,000)
on the frequency- of waves (Helix lactea ) .
No. of
A y "n p "P "1 TTi PTi "h
Frequency of waves
•r)c.-p minntp'
Control
Frequency of waves
•np*p Trni'Tii'i'hp
III JUXILA.
After dose of
adrenalin 1:10,000
2
66.0
67.8
3
76.0
67.2
7
85.4
80.1
9
79.2
77.4
10
64.6
88.2
13
74.3
60.7
14
79.2
. 72.0
TABLE Xd
Effect of doses of adrenalin chloride (1:10,000)
on the frequency of waves (Helix pomatia )
Ko. of Frequency of waves Frequency of waves
experiment per minute per minute
Control After dose of
adrenalin 1:10,000
60 96.0 92.0
62 103.1 81.3
66 95.2 92.6
69 60.0 87.0
70 62.2 94.0
75 71.9 62.0
72.
TABLE Xe
Effect of. doses of adrenalin chloride (1:10,000)
on the velocity of a single v/ave (Helix lac tea ) .
No. of
exper iaient
Velocity of a sinp:le
wave in rmn. per
minute
Control
Velocity of a single
wave in mm. per
minute
>-.fter dose of
adrenalin 1:10,000
2
2576
2682
3
3186
3060
7
5381
4200
9
3334
4704
10
4660
4358
13
6080
• 5250
14
3908
3510
TABLE Xf
Effect of doses of adrenalin chloride (1:10,000)
on the velocity of a single wave (Helix pomatia ) .
i:o. of
experiment
Velocity of a single
wave in mm. per
minute
Control
Velocity of a single
wave in mm. per
m.inute
After dose of
adrenalin 1:10,000
60
62
66
69
70
95
6920
7830
6010
5770
6280
6350
5400
5640
4950
8800
6460
5214
73
TABLE Xg
Effect of doses of adrenalin cnloride (1:10,000)
on the advance per wave (Helix lactea).
No. of
experiment
Advance per
wave in mm.
Control
advance per
wave in mm.
After dose of adrenalin
1:10,000
2
0.33
0.29
0.38
0.34
7
0.34
0.36
9
0.46
0.36
10
0.46
0.29
13
0.33
0.42
14
0.40
0.34
TABLE Xh
Effect of doses of adrenalin chloride (1:10,000)
on the advance per v/ave (Helix pomatia ) .
No. of
experiment
Advance per
wave in nim.
Control
60
52
66
69
70
75
0.96
0.45
0.19
0.78
0.34
0.45
Advance per
wave in !nm.
After dose of adrenalin
1: 10,000
0.24
0.28
0.38
0.50
0.31
0.32
74.
TABLE XI
Summary of effect of various concentrations of sdrenalin
on factors involved in locomotion.
Velocity of
Dose
1 of
adrenalin
!Number of cases
of increase
progression
or decrease from
normal
Plus
iviinus
0. 2
c c .
1: 10,000
1
6
0.3
cc .
1: 10,000
1
5
0.2
c c .
1 : 20 . 000
1
6
0.2
cc .
1:40, 000
2
4
0.2
cc .
a.: 80,000
3
13
0.2
cc .
1: 120,000
8
1
r I tJ u Lit? lio y ux
L> •
p
o
W ci V O O
O U •
p
0.2
cc.
1: 20,000
2
5
0.2
cc .
1: 40,000
1
5
0.2
cc .
1:80,000
2
14
0.2
cc .
1: 120,000
6
3
Vp 1 or i fv o f*
0.2
c c •
1: 10 .000
4
9
i3 J- t J. ti, JL w <V CL V ^
^ .
1*10 000
p
4
0.2
CC.
1:20,000
2
5
0.2
cc .
1:40,000
1
5
0.2
cc .
1 : oO , 000
14
0.2
cc .
1: 120,000
6
3
advance per
0.2
cc .
1: 10,000
2
5
wav e
0.3
cc .
1: 10,000
1
5
0.2
cc .
1:20,000
2
5
0.2
cc .
1: 40,000
1
5
0.2
cc .
1: 80,000
4
11
0.2
cc .
1: 120,000
5
4
75.
Recapitulation of TABLE XI
1: 10
,000
1:20
,000
1: 40
,000
1:80,000
1:120,000
+
+
+
+
+
Velocity of
progression
2
11
1
6
2
4
8 1
Frequency
of waves
4
9
2
5
1
5
7 9
8 1
Velocity of
a single
wave
4
9
2
5
1
5
2 14
^ IV
Advance
per wave
3
10
2
5
1
5
4 11
5 4
TABLE XIa
Effect of doses of adrenalin chloride (1:100,000)
on the velocity of proscression, frecuency of v/aves, velocity
of a single wave and advance per wave.
In U . O I
vexoGiuy QI
Velocity of progression
experiment
progression
in mm. per minute
in mm. per minute
After dose of adrenalin
Control
I : 100, 000
o
c,
oo • c,
45. 1
3
32.3
69. 1
18
68.0
80.8
21
42.0
40.4
28
81.3
47.1
Frequency of waves
Frequency of waves
per minute
per minute
After dose of adrenalin
Control
1: 100,000
2
95.2
90.1
3
78.1
121.0
18
64.5
95.5
21
71.1
109.3
28
99.0
83.0
Velocity of a single Velocity of a single
?/ave in min.per minute ' .^'ave in mm. per minute
after dose of adrenalin
Control
1; 100,000
2
4500
5120
3
5150
5500
18
5340
4420
21
3600
4500
28
6220
5930
Advance per wave
advance per wave
in mm.
in nun.
j.fter dose of adrenalin
Control
1: 100,000
2
0.40
0.46
3
0.41
0.48
18
1.18
0.65
21
0.63
0.73
28
0.61
0.47
ill
77
In order to determine whether the above results v/ere
due to the action of adrenalin and not to the chlorotone
present in the adrenalin chloride solution or a change in
Hydrogen ion concentration, dry suprarenal extract (Parke,
Davis & Co.) was nade up with Ringer's solution, A series
of experiments was carried out under the same conditions
as when adrenalin chloride was used. TablesXIIa through
Xllh give the data obtained when adrenalin chloride
1:80,000 and suprarenal extract 1:80,000 were used respec-
tively, and Tables Xllla through Xlllh give the data for
experiments when adrenalin chloride 1:120,000 and supra-
renal extract 1:120,000 v/ere used. A summary of this
comparison is given in Table XIV. Reference to this table
shows that results obtained from, suprarenal extract were
the same as for adrenalin chloride solution, except that
a greater depressant action was observed on the frequency
of waves with suprarenal extract of doses of 1:80,000.
The action of doses of 1:120,000 adrenalin chloride and
1:120,000 suprarenal extract was to cause either an increase
or decrease of the advance per wave.
Injections of adrenalin of any concentration within
the limits used, i.e., 1:10,000 to 1:120,000 during ver-
tical creepinp: did not usually affect the number of v/aves
present at one time on the foot. If there was a change
in the number it was by having one more wave present after
the injection of adrenalin. It was also consistently ob-
served after injection of adrenalin of either strong or
78
TABLE Xlla
Effect of doses of adrenalin chloride (1:80,000)
on the velocity of progression.
Ko. of velocity of progression Velocity of progression
experiment in ram. per minute in ram. per minute
i-fter dose of adrenalin
Control
1:80,000
8
23.8
14.0
10
64.5
35.0
11
35.0
16.0
,12
34.2
20.6
13
16.1
15.9
14
17.2
16.0
15
41.1
36.6
16
25.6
33.8
17
34.9
30.8
18
25.6
33.8
19
39.6
29.7
20
34.9
30.8
21
41.0
37.2
22
49.2
36.7
23
46.1
28.1
24
25.3
33.4
25
53.6
58.0
79.
TABLE XI lb
Effect of doses of adrenalin chloride (1:80,000)
on the freauency of v/aves.
^o, of Frequency of waves Frequency of waves
experiment per minute per minute
i-^fter dose of
Control adrenalin 1 ? P,f
8
75.5
• 71.4
10
135.0
87.6
11
87.0
83.0
12
62.8
63.0
13
62.5
61.5
14
69.6
60.1
15
82.1
61.0
16
59.0
72.0
17
70.8
106.6
18
59.0
72.0
19
74.3
65.5
21
72.5
60.0
22
104.0
72.4
23
75.4
78.0
24
61.5
70.0
25
90.7
94.1
80.
TABLE XIIc
Effect of doses of adrenalin chloride (1:80,000)
on the velocity of a single wave.
of
Velocity of a
Velocity of a
iriment
single wave in
single wave in
mm. per minute
mm. per minute
After dose of
Control
adrenalin 1:30,000
8
4600
3900
10
5160
4416
11
4800
4700
12
6300
5430
13
5620
5340
14
4450
3300
15
6230
5980
16
6250
5680
17
5060
4740
18
6250
5680
19
5700
4580
21
6600
7000
22
5340
4820
23
6080
5900
24
4150
4120
25
5580
5680
81.
TABLE Xlld
Effect of doses of adrenalin chloride (1:80,000)
on the advance per v/ave.
No. of
experiment
Advance per
wave in ram.
Control
Advance per
wave in mm.
After dose of adrenalin
1:80,000
8
0.31
■ 0.17
10
0.48
0.43
11
0.42
0.20
12
0.55
0.23
13
0.24
0.24
14
0.24
0.29
15
0.44
0.51
16
0.44
0.46
17
0.49
0.29
18
0.44
0.46
19
0.52
0.36
21
0.58
0.41
22
0.47
0.45
23
0.62
0.41
24
0.41
0.48
25
0.70
0.60
82.
TABLE Xlle
Effect of doses of suprarenal extract (1:80,000)
on the velocity of progression.
"No o"^
experiment
progression in
mm. per minute
Control
Vp"! op "i "h V nf*
progression in
mm., per minute
After dose of suprarenal
extract -1:80,000
124
26.4
28.5
136'
39.4
43.0
138
26.7
18.2
139
30.7
17.3
145
38.5
19.0
146
52.0
13.1
147
36.1
12.8
150
23.9
26.6
151
41.1
13.6
155
45.9
33.0
156
17.7
14.8
83.
TABLE Xllf
Effect of doses of suprarenal extract (1:80,000)
on the frequency of waves.
^o. of Frequency of waves Frequency of waves
experiment per minute per minute
Control
renal extract 1
124
79.1
78.0
136
76.6
61.2
138
82.3
79.1
139
52.8
66.1
145
64.6
58.6
146
92.6
90. 5
147
74.2
66.5
150
59.7
63.5
151
68.7
69.8
155
106.0
76.6
156
67.7
53.3
84.
TABLE XI Ig
Effect of doses of suprarenal extract (1:80,000)
on the velocity of a single wave.
No. of
Velocitv nf n
vexoGiLy 01 a
experiment
single wave in
single wave in
ram. per minute
mm. per minute
Control
After dose of suprarenal
extract r:80,000'
124
4010
4260
136
5100
5200
138
4360
4380
139
5500
4400
145
6040
5700
146
6216
5400
147
4600
3600
150
4400
3930
151
4900
3200
155
4520
4460
156
4550
3510
85.
TABLE Xllh
Effect of doses of suprarenal extract (1:80,000)
on the advance oer v/ave.
No. of Advance per wave .^-dvanct; per v;ave
experiment in mm, in mm.
i-.fter dose of supra-
Control renal extract 1;80,000
124
0.34
- 0.39
136
0.60
0.68
138
0.33
0.18
139
0.49
0.22
145
0.60
0.36
146
0.75
0.14
147
0.48
0.16
150
0.40
0.25
151
0.68
0.20
155
0. 56
0.36
156
0.26
0.29
86.
TABLE Xllla
Effect of doses of adrenalin chloride (1:120,000)
on the velocity: of progression.
Ko, of
experiment
Velocity of
progression in
mm. per minute
Control
Velocity of
progression in
mm. per minute
After dose of
adrenalin 1:120,000
7 ,
14.6
19.6
10
10.7
37.1
11
33.0
50.5
12
43.0
37.0
13
37.0
42.6
14
26.1
38. 1
21
14.3
19.0
25
10.2
12.8
26
26.0
38.1
87
TABLE XII lb
Effect of doses of adrenalin chloride (1:120,000)
on the frequency of waves.
^o* Frequency of v/aves Frequency of waves
experiment per minute per ?-ainute
After dose of
Control adrenalin 1:120^000
7 59.4 80.0
10 71.1 84.8
11 VI. 3 97.3
12- 68.1 127.0
13 126.6 91.2
14 70.0 100.0
21 58.6 84.7
25 52.3 62.0
26 70.6 100.1
88.
TABLE XIIIc
Effect of doses of adrenalin chloride (1:120,000)
on the velocity of a single wave.
of Velocity of a Velocity of a
experiment single wave in single wave in
mm. per minute mra.^'per minute
After dose of adrenalin
Control
■ 1:120,
7
5370
5730
10
4100
4600
11
6400
7030
12
6300
7990
13
7250
5850
14
5300
4700
21
3704
4880
25
3220
3760
26
5300
4720
89.
TABLE XI I Id
Effect of doses of adrenalin chloride (1:120,000)
on the advance per wave.
1:0. of
r-dvance per
wave .-.dvance per "vave
experiment
in mm.
in mm.
After dose of adrenalin
Control
1: 120,000
7
0,25
0.23
10
0.16
0.43
11
0.52
0.50
12
0.58
0.30
13
0.51
0.45
14
0.23
0.29
21
0.23
0.19
25
0. 18
0.16
26
0.23
0.29
90.
TABLE XI He
Effect of doses of suprarenal extract (1:120,000)
on the velocity of pror3:ression.
No. of
experiment
Velocity of
progression in
mm. per minute
Control
Velocity of
progression in
mm. per minute
After do3e of suprarenal
extract 1:120,000
36
47.1
49.2
38
37.2
52.6
40
23.6
54.2
42
23.4
48.6
44
25.2
48.2
45
50.1
65.1
91.
TABLE Xlllf
Effect of doses of suprarenal extract (1:120,000)
on the freouency of waves.
Ko. of Frequency of Frequency of waves
experiment waves per minute per minute
After dose of supra-
Control renal extract 1:120,000
36 100.0 102.0
38 71.2 110.7
40 65.5 89.8
42 79.7 88.4
44 63.4 93.0
45 82.3 112.1
92.
TABLE Xlllg
Lffect of
doses of suprarenal
extract (1:120,000)
on the velocity
of a single wave.
Ko. of
experiment
Velocity of a Velocity of a
single wave in single wave in
mm. per minute mm. per minute
After dose of supra-
Control renal extract 1:120,000
56
5604
6055
58
4540
5700
40
5840
5420
42
4660
4900
44
57 50
4890
45
4250
5020
93.
TABLE Xlllh
Effect of doses of suprarenal extract (1:120,000)
on the advance per wave.
Ko. of
Advance per
v/ave Advance per 'Nave
experiment
in ram.
in mm.
After dose of supra-
Control
renal extract 1:120,000
36
0.47
0.48
38
0.52
0.47
40
0.34
0.26
42
0.54
0.27
44
0.29
0.39
45
0.33
0.38
94.
TABLE XIV
Effect of doses 1:80,000 Effect of doses 1:120,000
Adrenalin Suprarenal adrenalin Suprarenal
chloride extract chloride extract
Plus Minus Plus i.Iinus Plus i.Iinus Plus l.Iinus
Velocity of
progression 3 13 38 81 60
Frequency of
waves 7
8
8
0
Velocity of
a single
wave
14
8
0
Advance
per v/ave
4 11
3 8
5 4 3 3
95.
weak concentration during vertical creeping that the ten-
dency to orient had been greatly lessened. previously
mentioned orientation and locomotion in Helix are closely
allied (Grozier and Navez, 1930) and further work is neces-
sary to disentangle the neuromuscular control of these
mechanisms •
Since there is no sympathetic nervous system in Helix
the usual explanation concerning the point of action of
adrenalin cannot be given. Adrenalin affects different
types of muscle differently. In structure the muscles of
the foot closely resemble other invertebrate muscle (Mendel
and Bradley). Elliott (1905) stated that adrenalin does
not excite the muscle fiber directly but acts on a substanc
at the m.yoneural junction. He stated that all other sub-
stances but adrenalin evoke, from, the group of plain m.uscle
tissues when stimulatin'i them directly, a reaction differin
only in degree and not in kind in each tissue. It is the
peculiarity of adrenalin to cause sharp contraction in one
and relaxation in the other. The cause is the same, the
effects different. Therefore the reacting substance must
be different and he has decided that since mechanical and
chemical stimuli do not point to marked intrinsic differ-
ences in the plain muscle fibers they do reveal differences
in the "nerve endings" and this difference is inherent in
the myoneural junction. Langley (1905-06) agreed with
Elliott as to the place of action of adrenalin but his
interpretation as to its mode of action differs from that
of Elliott. For Langley the dissimilarit-' of the sjTiaptic
96.
substance must be due to the intrinsic differences in
the cells in which the nerve fiber ends rather than to
intrinsic differences in the nerve fibers belonging to
any one system.
Gruber (1924) noted that adrenalin markedly shor-
tened the latent period and the duration of contraction
of a skeletal muscular contraction ?.nd induced a greater
shortening of the muscle v/hen stimulated, due possibly to
increasing the irritability or liberating more available
energy or actinpr catalytically on muscle metabolism.
Cannon (1929) considered that tiie effect of adrenalin is
on the muscle substance. Lutz (1930) interpreted the
inhibitory action of adrenalin on the heart of elasmobranch
fishes as the response of an unbalanced parasympathetic
mechanism in an organ lacking a sympathetic accelerator
innervation, and believed the action of adrenalin to be
on the nerve endings.
Cannon and 3acq (1931) have reported an adrenalin-
like substance which they considered a hormone "as it acts
in the same direction as sympathetic im.pulses and in their
absence is capable of bringing denervated organs into con-
formity with tnose v/hich are innervated, so that the caar-
acteristic unified response occurs." because this sub-
stance is derived from, stnactures under sympathetic control,
when they are influenced by s^nnpa thetic impulses, these
investigators suggested calling the substance sympathin,
but stated that it is only a provisional nam.e, for as
knowledge of its character increases it may prove really
to be adrenalin, developed for local action in smooth
viuscle. Wyman and Lutz (1932) have stated that further
wor:: is necessary to locate the region of the action of
adrenalin in Squalus acanthias .
Unpublished work (1933) of Cannon shov/s that sympathin
is not the same as adrenalin. Sympathin may be two sub-
stances, one excitatory, the other im.x jpy, called
E-sympathin and I-sympathin respectively. Both substances
are produced by the nerve endings, ks yet the exact
detailed interaction of these substances has not been
worked out.
From the results in this investigation it would appear
that the action of adrenalin is probably on the muscle
fibers rather than the nervous elements, .^s to its mode
of action on the muscle fibers it must be taken into
consideration that adrenalin is not natural to molluscs.
As far as can be determined in the literature the only
mention of this substance or a related one, was .nade by
Roaf and Nierenstein (1907) who stated that there is a
substance in the hypobranchial gland of Pii.ypura papillus
which is chemically and physiolo.:';ically allied to adrenalin.
Therefore adrenalin when injected into these animals may
act as a toxic substance, weak doses exerting a stimulating
effect and strone; doses an inhibitory one. On the other
hand the excitatory effects produced by weak doses of
adrenalin may be due to the fact that it improves the
98.
condition of vertebrate skeletal muscle and also is gen-
erally stimulating to invertebrate muscle. This improved
condition of the muscle may account for the increase in
velocity of a single wave, the velocity of prof^ression
and the frequency of waves.
The possibility that adrenalin increases, the irrita-
bility of muscle may account .also for the increase of tnese
activities when. weak doses are used. It is conceivable
that if adrenalin acts on the rayo-neural substance T-ich
substance may have different degrees of stajility. i'.ie
strena;th of the dose of adrenalin may cause changes in
this unstable substance, which in turn causes ~reater or
less activity of the factors involved in locomotion.
The proportionality between the rate of creeping and
the velocity with which a single wave traverses the foot,
or between the freo_uency of .the pedal waves and the
velocity of progression is not altered in Helix by the
injection of .adrenalin: these facts further support the
idea of m.uscle as an "independent effector".
G. Effect of strychnine sulphate on locomotion.
Crozier and Federiehi (1924-25a) have pointed out
from their work on the effect of temperature changes on
phototropic circus movements of Limax that "a certain
type of Dredict^bT lity in ani'^^-i --^ -"T-der 'nor'"'.?!'
natural conditions probably results from dynamic equilibrium
9Q.
thereby obtained between diverse '■^oo'--': r i?^ ^ ^ '^-^ ting for
effector control (in the present case, trie creeping
mechanisra and that for turninc^, in the ran^^e 14° - 460G.).
It follows that unravelling of the elements of conduct
necessitates experi;r.entation under diverse abnormal con-
ditions favoring individual mechanises of r'=^sPons°."
Th"3e investi£;ators have succeta-o^ 1 : ^ - -a- " :
^^t»..^.i — j.^
these two activities, , orientation and locomotion.
In the experiments on phototropic circus mover.-^nts of
^^^^^ as affected bj changes in temperature tnese authors
found that below 15° the amount of circling is determined
very largely by speed of creepin*?, v/hil^^ o>^ov9 15O the
pedal m-echanism is in secondary place, the turning
mechanism becoming the controlling element.
Another method of dissociatir ~ t-ese t^-':^ • -^chanisms
was by the injection of strychnine. This substance was
found to suppress the phototropic circus movements of
Limax msximus (Crozier and Federighi, I'^'^'—^d'r), but no
detailed information was obtained concerning the changes
in the creeping activity of the foot. Therefore the sub-
sequent experiments were performed to study the effect
of strycrjiine on the behavior of the pedal waves and to
compare them with the normal foot. Small doses of
strychnine of definite concentration disturb the functions
of synaptic nervous systems, while the same concentration
does not disturb the non-s7rnaptic net. By the use of
this substance Crozier and Arey (1919) found that in the
100
mo Husk Ghromodorls the general in te "lament rr-3 O "! ''^-o
outgrov/ths of the body depend upon a locally contained,
peripheral, non-synaptic net-work, but the central
nervous system is essentially synaptic. ';/:aile a large
proportion of raollusks have the entire nervous system of
the non-synaptic type, those v/hich show reversal of in-
hibition possess a synaptic nervo^is system.
The ef feet . produced by strychnine, i.e., "reversal
of inhibition" is interpreted in various ways by different
authorities. In the spinal cord of vertebrates it is
usuall3r supposed that tlie strychnine effect is due to the
abolition of the inhibitory component of normal coordina-
tion, so that the inhibitory effect is transposed into
an excitatory one.
. M. and time. Lapicque (1908) have concluded that the
effect of strychnine is to bring about a condition of
isochronism, i.e. , the chronaxie of the nerve equals that
of the muscle fiber and thus the excitation spreads
easily from one neurone to another.
It may be argued that the action of strychnine is
not due to true reversal but is merely due to a condition
of augmented central excitability in -.vhich the excitatory
effects produced by stimulation of a mixed efferent nerve
conceal the inhibitor:/ effects of the fibers of the sam.e
group, i.e., a homogeneous group of inhibitory impulses
is not converted into excitatory impulses by strychnine.
Magnus and Wolf (1913) and others have observed conditions
101.
which support this interpretation.
Bayliss (1918) has concluded from his experiments on
vascomotor reflexes that there may be two independent
synapses with which the firr] co'r.rror! notor path connects,
unequally sensitive to strychnine, or that the drug acts
on some intermediate synaptic membranes on the afferent
side, sjTiapses which are not part of the path comraon to
the different reflexes.
Bremer and Rylant's (1924, 1925) theory of "reversal"
by strychnine is in accordance with that of the Lapicques.
They believed that strychnine breaks down excitatory
barriers between neurones, thus allowing excitation to
spread freely from neurone to neurone throughout the
nervous system. It does this by equalizing the chronaxie
between various contiguous units which in the nor^.ril un-
poisoned nervous system are separated by a bari-ioi- of
heterochronism.
Fulton (1926) concluded that true reversal of inJiibi-
tion into excitation probably does not occur, ana the
apparent reversal is due to the stimulation of excitatorj^
fibers in a mixed nerve, which, cvin'-.- to the increase of
excitability produced by strychnine, conceals tiie effect
of concomitant inhibition.
Various investigators have recently suggested that
too much importance has been given to the question of
chronaxie, and have questioned Lapicque's interpretations.
Rushton (1930) did not agree with his claim that the
102.
chronaxie of muscle is normally the same as that of the
nerve '.vhich supplies it. i-.e strongly supported ^.ous
(1907) who maintained that there are two excitable sub-
stances in muscle of which one has a very much lar!7er
chronaxie than nerve. F-urther, Kushton (1S.32) nas also
questioned Lapicque's terminology. Ke (Rushton) stated
that "since Lapicque wishes to restrict the n-'i'-rie 'chronaxie'
to 'true' chronaxie, it is important to nav^ a nev/ term
which can be applied to any strength duration curve 'true'
or 'false'. Lucas 'Excitation Time' 'vss ^ic^-r. 'jst
this sense and it is proposed for a-or. :ior. . ' ^r.afest
(1932) also stated that Lapicque's law of isocnronism is
not valid, at least for excitability of the -i---''^
fiber nerve-muscle complex. .-.s agreed w;izr. zr.e views neld
by Lucas and Rushton. Lapicque (1932) did not think that
Rushton's investigations ^v^^..^ ^^y. lio-ht on
this problem and r.e defenaea r.is position against Rushton.
It can be seen that too great importance has been placed
on the sub.iect of chronsxie before the various laws
claimed for it .lave oeen satisfactorily interpreted and
verified.
Although these various theoretical opinions concerning
reversal of inhibition exist, it is possible tr_pough pre-
cise m.easureraents to obtain data on the behavior of Helix
v/hen injected with strychnine. In this w^. - i-^ - be
possible to gain information about central nervous
activities which could not be obtained by other means.
103.
Although Grozier and Pederighi (1924-25b, 7-221-224)
were chiefly concerned with the effect of strychnine on
the phototropic movements of Limax maximus and not pri-
marily with locomotion, they state that strychnine does
not essentially affect locomotion. It was the purpose,
therefore, of this investigation to study the character-
istic effect of various amounts of str^/chnine on Helix
lac tea and Flelix pomatia during locomotion. As far as
can be determined no mention of work of this nature can
be found in the literature. A study was also made at the
same time of the effects of strychnine sulphate, 1 part
in 100, on the animals creeping in various planes, i.e.,
vertically upwards, horizontally (under surface) and
horizontally (upper surface).
A series of experiments was first perform.ed to study
the effect of strychnine sulphate in the concentration
of 1:1000 (1 part in 1000 parts of ninger's solution).
Measurements were obtained for the controls and for the
in.iected animals during vertical creepir.-. After 0.2 cc.
of strychnine sulphate was injected there was either
partial or complete retraction of the foot into the
shell. The time elapsing after injection before creeping
began again varied from fifteen minutes to one hour. It
was found that with strychnine sulphate, 1:1000, the
creeping was more or less regular and could be compared
to normal creeping. The specific results for Helix lactea
under these conditions are given in Table XV. Table XVa
104.
TABLE XVa
Effect of doses of str3'-chp-ine sulphate,
1:1000, on the velocity of progression (V), the
frequency of v/aves (F), the velocity of single
wave (v) and the advance per wave (A).
■^Plus Minus Cases unchanged
V 3 4 0
F 3 2 2
v 2 5 0
A 0 3-4
Total number of increases or decreases
from the normal animal.
105.
shows that the factors involved durinp- loco-otio- not
essentially altered.
Injections of this concentration did not seem to
affect the amplitude of the waves, as it -!-3 the same or
nearly the same as when the animal was creeping under
normal conditions (Table XV, advance per wave).
The effect of stronger concentrations of strychnine
were studied for nineteen animals. In the experiments
forty-eight series (7150 waves) were obtained for the
control animals and forty-four series (6550 waves) for
the injected ones v/hen 0.2 cc. strychnine sulphate,
1:100, was injected. Creeping began from fifteen minutes
to one hour after the injection. The data obtained from
these experiments are given in Table XVI. Reference to
Table XVIa shows that strychnine of this concentration
did not affect any of the factors involved in locomotion.
Strjrchnine of still greater concentration was used
(1:50) but no results could be obtained because after
injection when the animal began creeping, sometimes after
one or two hours, the creeping was irregular and it was
impossible to count the v/aves.
The specific effect of the gravitational pull on tne
normal and s trychninized animal during locomotion in three
different planes, i.e., vertical, horizontal (under sur-
face) &nd horizontal (upper surface) was analyzed. The
procedure was to place the animal on a horizontal glass
plate which rested on supports about six inches high
106.
TABLE XV.
The effect of strychnine sulphate (1:1000) on vertical
ascensions of Helix lactea."^
i\ 0 . 01
Velocity of
Frequency.''
Velocity of
iidvance
experiment
progression
of waves
sin2;le wave
per wave
in mm. per
per minute
in mjn. per
in m]'.
minute
minute
#28 (normal )
9 . 1
45.0
3650
0.20
ffeiy V U • D CC •
s tr ychnine
sulphate
1:1000)
10.0
52.0
7700
0.19
#30(Kormal)
11.2
52.0
7100
0.21
#31(0.5 CC.
strychnine
sulphate
1 : 1000 )
11.8
55.0
8300
0. 21
#32 (Hormal )
4.6
62. 5
6550
0.07
#33(0.5 CC.
"h"r*vp Vtp 1 n ft
sulphate
1:1000)
5.7
62.5
5800
0.09
#57 (Nor-iia 1 )
20. 6
100.0
4500
0.24
#58(0.3 CC.
sulphate
1:1000)
19.0
83.0
4100
0.14
#61 (r ormal )
23 . 1
oo . 0
4oUvJ
r\ or*
#52(0.3 CC.
strychnine
sulphate
1:1000)
8.5
71.0
4600
0.12
#82 (Normal)
16.8
83.0
4600
0.20
#83(0.3 cc.
strychnine
sulphate
1:1000)
14.2
100.0
4500
0.14
#84(norrnal )
27.0
100.0
7100
0.27
#85(0.3 CC.
strychnine
sulphate
1 : 1000 )
22.0
100.0
4100
0.22
Temperature limits were 16.6° to 21.1°G. but the temperature
was constant for each experiment.
107.
TABLE XVI
The effect of stryclinine sulphate (1:100) on
vertical ascensions of Helix lactea.
No. of
Velocity of
Frequency
Velocity of
^.dvance
exoer inpn t
o I wa V e s
a single wave
per v/£ve
in ram. per
per minute
in mm. per
in rnrn .
LliX ii u. 0 "
minute
4 0
60,6
81.2
3828
0.48
5 e
. 72.1
90.0
4069
0.54
6 0
42.0
72.6
3078
0.36
7 0
39.2
77.4
3772
0.42
8 0
64.8
67.8
3109
0.40
9 •
57.0
79. 2
19 0
57.3
24.9
78.1
0.34
20 A
s"^ n
oo . u
<cO . c.
74. 5
0.28
21 0
46.2
29.7
112.0
0.31
p
o^t . o
o D . y
OC . 1
0. 42
23 0
62.8
31.7
90.3
0.40
24 •
55.6
18.9
82.8
0.23
25 0
21.9
96.0
4950
0.29
2d e
19 . 4
70.0
4800
0.27
or? „
27 0
20 . 6
84.2
5150
0.24
28 e
21.3
56.9
4900
0.38
29 0
31.1
91.7
4660
0.31
30 •
31.3
97.0
4920
0.29
31 0
20.8
68.0
5900
0.29
o2 9
27.0
96.6
6175
0.29
33 0
25.0
89.2
5140
0.30
34 •
13.5
69.2
4420
0.20
o
o
= control
= injected 0.^1:^1.3] (0.2 cc . of st^^''?"o''inir ^ 9-1 f-p 1:100)
108
TABLE XVI (cont. )
No. of
Velocity of
in mm. per
minute
Frequency
u± waves
per minute
Velocity of
a single wave
in mm. per
minute
i.dva"'' '
per
in mm.
35 0
36 0
39.6
48.6 .
96.0
54.0
4300
5400
0.44
0.35
37 0
38 •
21,0
13.0
56.0
96.0
4420
0.24
0.18
39 0
40 •
28.8
32.1
88.0
86.9
5010
0.52
0.42
88 0
89 0
29.0
23.1
111.0
68.0
5400
3400
0.26
0.30
90 o
91 •
30.8
11.1
108.0
97.2
4600
4600
0.29
0.10
92 0
93 •
35.0
16.3
98.1
92.1
5500
5160
0.30
0.21
94 0
95 «
35.6
40.0
76.2
ei.o
3500
4000
0.13
0.16
96 o
97 •
13.4
25.0
72.2
98.3
4900
5400
0.19
0.26
o
= control
= injected animal (0.2 cc. of strychnine sulphate 1:100)
109.
TABLE XVIa
Effect of strychnine sulphate, 1:100, on the
velocity of progression (V), the f renu.eno^'- of waves (P)
the velocity of a single v/ave (v) anu L.ia advance per
wave (A).
■^Plus Minus Cases unchRriQ;ed
V 7 10 -2
F 9 10 0
V 9 9 1
A 7 8 4
^ Total number of increases or decreases from the
n or''^/-'' s ""^ i'" 1 •
110
(F±p^, 15). Under the glass plate a mirror was placed at
a slope of 45° on a block of wood for the purpose of study-
ing the waves when the snail was creeping on the upper
surface of a horizontal plane. The methoa or measurement
was the same as given above. Data were first obtained for
the normal animal when creeping in (1) a vertical plane,
(2) in a horizontal plane (under surface) and (3) Horizontal
plane (upper surface); after injection of strychinine sul-
phate measurements were obtained in the same plane as for
the normal animal. The actual scatter was plotted of the
various relationships and the slopes of the line were
compared.
Pigs. 16, 17 and 18 give typical results and show
that in all planes for the s trychninized animal a linear
relations'^ip exists between the velocity of a single v/ave
and tne velocity of progression, between the frequency
of waves and the rate of creeping and between the advance
per wave and the rate of creeping. A smaller change in
the slope of line is observed for the relationship between
the advance per wave and the velocity of progression
(Fig. 18) than is noted for the other two relations (Fig.
16 and Fig. 17).
The effect of the gravitational pull on the normal
animal during locomotion in the three different planes
was observed by comparing the slope of the lines obtained
for each relationship. The slope of the lines showing
the relationships occurring during vertical creeping was
111.
usually less than for the same relations durlnfr creeping
on the horizontal plane, upper or under surface, defer-
ence to Table XVII shows that the velocity of a single
wave, freouency of waves, the velocity of progression and
the advance per wave v/ere usually greater when the animal
was creeping in a horizontal plane (upper surface) and
creeping vertically upwards than when creeping in a hori-
zontal plane (under surface).
In the s trychninized animal not only the change due
to strychnine, out also the increase or decrease due to
changes in gravitational pull was determined. (Fig. 16,
17 and 18. ) i-xS the gravitational pull is decreased there
is usually an increase in the slope of the line describing
the various relationships when the animal was creeping
on the horizontal plane, upper surface. This increase
was less as the gravitational pull was increased (vertical
creeping and creeping on a horizontal under surface).
However, the results show that when an animal was creeping
in each of these two planes tnere was a greater alteration
of these factors than when creeping on a horizontal upper
surface (Table XVII).
The results in this investigation also show that as
the effect of gravity was lessened the factors involved
in locomotion were increased, for creeping was increased
in the horizontal plane (upper surface) as compared with
the other planes. Cole (1925-26) has stated that the
stimulus for geotropic orientation and locomotion is the
112.
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114.
tension of the body muscles produced by the downward pull
of gravity; this stimulus is received by the propriocep-
tors of the muscles. However, if this were the condition
we should not expect to get faster creeping when the pull
of gravity is lessened; consequently. Cole's theory about
locomotion, while it may hold true for orientation, does
not seem to sojve the problems involved in locomotion.
The fact that strychnine sulphate in the strongest
dose that could be used practically (1:100) did not-
essentially affect locomotion may be due to the fact that
invertebrates are relatively more insusceptible to
strychnine than vertebrates. Also if strychnine acts on
synapses, the lack of the usual excitation may be due to
a lack of synapses in the intrinsic mechanism of the foot.
Strychnine in this investigation did not abolish the
linear proportionality between the velocity of a single
wave and the velocity of progression, between the frequency
of waves and the velocity of progression and between the
advance per wave and the velocity of progression.
115,
Chapter VI.
Surnmary and Conclusions
1. The results of these experiments show that in
Helix lactea the speed of creeping vertically upward,
when not carrying loads, is directly proportional to the
velocity of a single pedal wave. The speed of creeping
is also directly proportional to the frequency of the
waves and to the advance per wave. The proportionality
factor is independent of the weight of the animal.
2. The proportionality between the rate of creeping
and the velocity with which a single wave traverses the
foot, or between the frequency of the pedal waves and
the velocity of progressibn, is not altered in normal
or in de-eyed Helix when lifting loads during vertical
creeping. Under these conditions the linear relation-
ship between the advance per wave and the rate of creeping
is not altered. The foot appears to be essentially an
independent effector, although under the control of central
impulses .
3. The detached foot was observed and kymograms
were obtained which showed that the rhythmical action of
the foot is greatly interfered with when it is not con-
nected with the pedal ganglia. The destruction of the
periodic recurrence of waves when the ■oedal 'mnp;lia are
separated from the foot suggests that tnis periodicity
is determined by the presence of these ganglia.
116
4. Injection of Ringer's solution or raechanical
stimulation of the body wall or anterior end of the foot
does not affect the factors involved in locomotion.
5. Adrenalin of various concentrations does not
alter the linear proportionality between the velocity of
progression and the velocity of a single wave, between
the frequency of waves and the velocity of progression
and between the advance per wave and the rate of creeping.
These are additional factors which establish tne idea of
the foot
"independent effector" o^/ muscle.
Adrenalin in concentrations of 1:120,000 (0.2 cc. or
0.3 cc. ) produces a stimulating effect on the velocity of
progression, frequency of waves, velocity of a single
wave and the advance per wave.
An inhibitory effect of adrenalin on invertebrate
tissue not previously reported is given. ^-.drenalin in
concentrations of 1:10,000, 1:20,000, 1:40,000 and 1:80,000
produces a depressant action on the velocity of progression,
frequency of waves, velocity of a single pedal wave, and
the advance per wave.
The region of action of the adrenalin is discussed.
6. Strychnine sulphate (1 in 100, 1 in 1000) does
not bring about reversal of inhibition in Helix. It does
not abolish the linear proportionality between the velocity
of a single wave and the velocity of progression, between
the frequency of waves and the velocity of progression
and between the advance per wave and the velocity of Pro-
gression. These facts are additional evidence that the
117.
foot is essentially an "independent effector".
The effect of , gravitational pull on the normal and
s tryclininized animal during locomotion in various planes,
i.e., vertically upwards and horizontally (upper surface)
is usually to increase the velocity of a single wave,
the frequency of waves, the velocity of progression and
the advance per wave, while during creeping in horizontal
plane (under surface) a decrease in each activity occurs.
7. The purpose of this inves tig;ation was to estab-
lish if possible the factors in control of locomotion in
Helix. It has been shown that the foot of Helix is pri-
marily an independent effector and is secondarily under
the control of central impulses. In other words, the
intrinsic neuro-muscular mechanism of the foot is the
primary factor in locomotion.
118.
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ten Gate, J. Quelques recherches sur la locomotion des
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Grozier, W.J. and Navez, k.E. The geotropic orientation
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ti
Jordan, H. and Hirsch, G.Ghr. Ubungen aus der Vergl. Physiol.
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121
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123.
Index of Figures
Fig. Al. Diagrams showing: Page 127
a. Helix poniatia, dorsal vie'v
b. " " ventral view
0. " lactea, dorsal view
d. " " ventral view
A2. Diagram showing nerve-net in foot of Helix 128
A3. Diagram of Helix showing arranp-ement of ganglia
(cerebral,' pedal and visceral) 129
A4. Schematic representation of the musculature
of the foot of Helix 130
1. Diagram of the method of measurement for
vertical creeping 131
2. The linear relationship between the velocity
of a single pedal wave and the rate of creep-
ing during vertical ascensions without load 132
3«. The relation between the velocity of progres-
sion and the frequency of waves for vertical
creeping without load 133
4. Tlie relation between the velocity of progres-
sion and the advance per wave for locomotion
in a vertical plane without load 134
4a. Comparative graph showing the relation between
the velocity of progression and the velocity
of a single wave without added load 135
4b. Comparative graph showing relation between
the velocity of progression and the frequency
of waves without added load 136
4c. Comparative graph showing relation between
velocity of creeping and the advance per wave
without added load 137
4d. The relation between speed of creeping and
velocity of a single wave without added load
in Helix pomatia . 138
4e. The relation between velocity of progression
and the freouency of waves without added load
in Helix poiaatia. 139
4f • Relation between the velocity of progression
and the advance per wave without added load
in Helix Domatia. 140
124
Page
Fig. 5. The effect of added loads on the relation
between frequency of waves and the rate of
creeping 141
5a. Actual scatter obtained frotn series of ver-
tical ascensions with and without loads showing
the relation between the velocity of progression
and the frequency of waves 142
5b. Relation between the rate of creeping and the
frequency of waves with added loads during
vertical ascensions for animal No. 10 143
6. The relation between tension (added loads) and
the rate of creeping - 144
7. The probable error for ti:ie rate of creeping of
animal No. 6 as a function of tension (added
loads ) 145
8. Comparison of the normal and de-eyed animal
showing the relation between the velocity of a
single wave and the rate of creeping 146
8a. iictual scatter obtained from series of vertical
ascensions without load for normal and de-eyed
animal showing the relation between the velocity
of the pedal waves and the rate of creeping 147
9. The relation between the frequency of waves and
the rate of creeping for de-eyed Helix lactea 148
9a. Actual scatter obtained from a series of vertical
ascensions without load for normal and de-eyed
animal showing the relation between velocity
of progression and the frequency of waves 149
10. The relation between the advance per wave and
the rate of creeping in the normal and de-eyed
animal 150
10a. Actual scatter obtained showing the relation be-
tween the velocity of progression and the advance
per wave for animal No. 3 v/ith and without eyes
carrying no load 151
10b. actual scatter showing relation between the
velocity of pedal waves and the rate of creep-
ing for normal and de-eyed animals with added
load of 0.7 gram 152
10c. iiCtual scatter showing relation between the
velocity of progression and the frequency of
waves for normal and de-eyed animal with added
load of 0.7 gram 153
125.
Page
Pig.lOd. nctual scatter showing relation between the
velocity of progression and the advance per
wave for normal and de-eyed animal with added
load of 0.7 gram 154
lOe. Comparative graph showing relation between
velocity of a single wave and the rate of
creeping for de-eyed animal carrying varying
loads 155
11. Diagram of the arrangement for recording the
behavior of the detached foot 156
11a. Kymograph records of the responses of the
and b. detached foot 157
12. The effect of adrenalin on the relation between
the velocity of a single pedal wave and the
rate of creeping 158
13. The effect of adrenalin on the relation between
the frequency of waves and the rate of creeping 159
14. The advance per wave as a function of the
rate of creeping for the normal and injected
animal (adrenalin) 160
15. Diagram showing the method of measurement
for creeping in a horizontal plane (upper
surface) 161
16. The effect of strychnine on the relation
between the velocity of a single pedal wave
and the velocity of progression 162
a. Creeping in vertical plane
b. " " horizontal plane (under
surface )
c. " " " " (upper
surface )
17. Comparison of the normal and injected
animal (strychnine sulphate) showing the
relation between the frequency of waves and
the velocity of progression 163
a. Creeping in vertical plane
b. " " horizontal plane (under
surface )
c. " " " » (upper
surface )
(
Comparison of the normal and injected
animal (strychnine sulphate) showing the
relation between the advance per wave and
the velocity of progression
a. Creeping in vertical plane
^» " " horizontal plane (under
surface )
c. " " " " (upper
surface )
Figure A 1
Diagrams showing:
^* Helix po!;iatia , dorsal view
b. Helix pomatia, ventral view
c. Plelix lactea , dorsal view
d. Helix lactea, ventral view
FIGURE A I
Figure A 2 ♦
Locomotor nerves of Helix pomatia.
(a) Kerve net in foot.
(b) Central part of nerve system.
1. Ccrebro-pleural connection,
cut.
2. Cerebro-pedal connection.
3. Radiating nerves of the
pedal ganglion.
4. Cerebral ganglion.
5. Pedal ganglion.
(From Jordan, p. 225, after E. S.chmalz,
1914. )
FIGURE Ai
I
Figure A 3
Diagrams showing arrangement of cerebral
ganglion (c), pedal ganglia (P) and visceral
ganglia (v).
(Modified after Richard Hertwig, "A
Hanual of Zoology" by Richard Hertwig, trans-
lated and edited by J. S. Kingsley, p. 552.)
I
Figure A 4
Schematic representation of the musculature
of the foot of Helix. The longitudinal muscle
fibers are red, the dorso-ventral black. Pig. 5
is a horizontal, Fig. 6 a vertical longitudinal
section. Of the retractor bundles, which are
to be considered in Fig. 5 as spread out,
only three are represented, rl, r2 and r3.
(Taken from Zeitscnrif t f^r wissenschaf tliche
Zoologie. 30er Band. Tafel acht. Supplement,
Erstes Heft. Leipzig 1878.)
I
Figure 2
The velocity of a single wave is directly
proportional to the velocity of progression
during vertical ascensions without load.
The average deviation of the mean for each
point plotted for animal Ho. 7 (Kelix lactea)
is as follov/s:
Point 1 1*0
'12 0.25
" 3 0.53
"4 2.9
"5 1.5
Aniraal V/eight of Slope of
animal line
No. 1 6.5 gm. 0.2
II 4 8.6 " 0.36
n 7 7.9 " 0.34
f
A.D.=I
<
O
-J
—
3
O
f
I.
i
o
O
LU
to
1 1 1
\ 1
0
n
ve
1 •
o
<^
1 1 ! rv
II ^
-e-
to
O
o
_J
1 1 1
>
1
1
1 \
L
1
\ o
7)
QO
CO
lO
JO JiVd
/wiJ V vji \jiu_]_iaj
Figure 3
The frequency of the pedal waves is
directly proportional to the velocity of
progression, during vertical ascensions
without load. This proportionality holds
regardles^^ of the weif?ht of the animal. (Helix
lactea). (Tahle III).
Animal Weight of Slope of
animal line
IJo. 1 6.5 gm. 0.2
n 7 7.9 " 0.18
" 10 3.8 " -0.2
Figure 4
The advance per wave is proportional to
the velocity of progression in vertical creeping
without loads. This relationship holds regard-
less of the weight of the animal (Helix lactea).
(Table III).
Animal Weight of Slope of
animal
line
No. 4
" 10
8.6 gm.
0.3
0.3
0.3
Figure 4A
This comparative r;raph shows that the
velocity of progression without loads during
vertical ascension is directly proportional
.to the velocity with which a single wave tra-
verses the foot and that it is independent
of the weight of the animal (Helix lactea).
Animal Weight of Slope of
animal
line
No. 1
" 3
t» 4
" 5
" 6
w Y
" 8
" 10
" 11
6.5 gi
7.7 ^
8.6 "
7.1 "
6.7 "
7.9 "
8.5 "
3.8 "
5.0 "
:m.
0.2
0.52
0.36
0.48
0.82
0.34
0.52
0.54
0.24
Figure 4B
The frequency of the pedal waves is
directly proportional to the velocity of
progression during vertical ascensions
without loads. This proportionality holds
regardless of the weight of the animal
(Helix lactea ) .
1 1
z
(0 u.
irj 09 lo t>»>et^
1°
z
<0 =
I I
I I
I 1
T T
o
u
>"
e3
00
Q-
LJ
Ld
cr
o
u
h-
cr
CQ
cr
Z)
^ — O O) 00
Figure 4C
In the analysis of nine animals (Helix
lactea) for the relationship between the
advance per wave and the velocity of pro-
gression the advance per wave is propor-
tional to the velocity of progression in
vertical upward creeping without loads, and
this relationship holds regardless of the
weight of the animal (Table III).
Figure 4D
The rate of creeping during vertical
ascensions is directly proportional to the
velocity with which a single wave courses
over the foot v/hen carrying no load. This
relationship is illustrated for animals
Number 41, 44 and 46. (Helix pomatia. )
-Figure 4E
The rate of creeping is directly
proportional to the frequency of pedal
waves during vertical ascensions without
loads. This relationship is illustrated
for animals I'lumber 41, 44 and 46. (Helix
por.iatia . )
Figure 4F
The velocity of progression is pro-
portional to the advance per wave during
vertical ascensions without loads. This
is illustrated for animals Number 41, 45
and 46. (Helix pomatia. )
Figure 5
In Helix lactea the linear propor-
tionality between rate of creeping (V) and
frequency of the pedal waves is not altered
when lifting added loads during vertical
creeping. This graph shows the individual
effect of adding varying loads on animal
Number 7.
Slope of line
Without load
0.20
With load of 0.7 gm.
1.2 "
2.0 "
2.5 "
3.0 "
0,20
0.22
0.18
0.42
0.20
0.22
0.22
I
II
I
Figure 5A
The actual scatter obtained from
direct observations for a series of runs
with and without loads during vertical
ascensions. The relationship between the
velocity of progression and the frequency
of waves has been plotted for animal
IJumber 7. (Helix lactea. )
©
o
o
o o
^ o
© o
0>
o o ■ f) o
O * ^ <] «o
^ <, Sp ^
z uJ X O ©0
< to H © ©
tc < ^ ^ lo
UJ li- _ '©
|_C/) ^' J ©
< z F m
O O I- 0 o
_jf ^ + ©
© <
O CD 4.
< O o
<o<C
^ m
5tt>-Cv20lOOQDro CD
"^O ©©□><•■ +
CO
oj — O <i> CO r>-
Hi
ii
Figure 5B
This comparative graph shov/s the rela-
tion between the rate of creeping and the
frequency of waves for animal Number 10
(Kelix lactea ) with added loads during
vertical ascensions.
Added load
0.7 gm.
1.2 "
2.0
2.5 "
3.0
3.8
4.8 "
Slope of line
0.18
0.16
0.16
0.14
0.18
0.14
0.28 exhaustion
O CT) 00
1 '
Figure 6
The relation of tension to the rate of
creeping is shown from data obtained from ani-
mals number 1, 3/4, 6, 7 and 8 during upward
vertical creeping (Kelix lactea). The tension
is represented by the addition of varying
loads, i^., 0.7, 1.2, 2.0, 2.5, 3.0, 3.8,
and 4.3 grams. With a great many animals
the effect of a load of 2.5 gm. was a decided
decrease in the rate of creeping. Usually a
slight increase occurred with the addition of
3.0 grams.
I
Figure 7
Figure 7 shows the probable error
(calculated according to Bessel's formula)
for the rate of creeping with added loads.
Ci>-
_CvJ
o
(f)
I I L_
(O rj-
9Nld33H3 JO
to
Figure 8
The removal of the eyes does not
alter the law of linear proportionality
between the velocity of a single wave
and the rate of creeping without loads.
(Helix lactea. )
Figure 8A
The actual scatter is shovm of data
obtained from direct observations of the
normal and de-eyed animal (Helix lactea) .
The relation between the velocity of the
pedal waves and the rate of creeping has
been plotted for animal Ko. 3 during
vertical ascensions without load.
CO
*
_J
a
u
I-
1-
<i:
O
_i
ZD
I-
o
o
mm
o
o
o
llJ ^
V 111
OJ >
UJ
O
Q
O
h
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I
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o
o
o
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o
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o
8
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I
D
Q.
Q.
4j_
O
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o
in LlI
CO >
QO
Uj
q:
L.
00 <D -sh
I
Figure 9
The linear relationship between rate
of creeping and frequency of waves is not
altered in the de-eyed animals without
load (Helix lactea).
Figure 9 A
The actual scatter is shown of data
obtained from direct observations of the
normal and de-eyed animal (Helix lactea ) «
The relation between the velocity of
progression and the frequency of waves
has been plotted for animal llo, 3 during
vertical ascensions without load.
zr-ir
m
o
o
8.
CO
hi
Q
• o
Q
<C
O
-J
h
O
X
h
IjJ
1-
O
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13
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o
o
o
o
in
o CT) 00 vD
(0
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>
o
o
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CD
U
cr
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Figure 10
The relationship between the rate of
creeping and the advance per wave is not
altered in the de-eyed animal without load
(Helix lactea).
|A||A|'3AVM H3d B^NVAQV
I
I
Figure lOA
The actual scatter is shown of data
obtained from direct observations of the
norrnal and de-eyed animal (Helix lactea ) .
The relation between the velocity of
progression and the advance per wave has
been plotted for animal No. 3 during
vertical ascensions without load.
o
o
o
o
CO
<
z
<
q:
hi
h-
I-
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o
(f)
-I
<
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<
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if)
t Id
• o
O
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8
o
O CD CO ^• vo m
Figure lOB
The actual scatter is shown of data
obtained from direct observations of the
normal and de-eyed animal (Helix lactea ) .
The relation between the velocity of the
pedal waves and the velocity of progression
has been plotted for animal No. 3 during
vertical ascensions with added load of 0.7
o
• ^ X UJ
• •
^ Q
• O
o
o
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o
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8
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— doo dbod
33«^w X'NOI9S3d90ad dO AllQ0n3A
Figure lOG
The actual scatter is shown of data
obtained fror.i direct observations of the normal
and de-eyed animal (Helix l&ctea ) « The rela-
tion between the velocity of progression and
the frequency of waves has been plotted for
animal Ho. 3 during vertical ascensions
v/ith added load of 0.7 gin.
t • •
o
o
8
X u
• o
u
U
CO
<
D
r
o
d
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t
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Figure lOD
The actual scatter is given of data
obtained from direct observations of the
normal and de-eyed animal (Helix lactea ) .
The relation between the velocity of
progression and the advance per wave has
been plotted for animal Ko. 3 during
vertical ascensions with added load of
0.7 gm.
o
o
0)
SQ
• o
o
o
•
• •
•
•
o
o
0
O 0
x:
cr
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-I
° 8
o o
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o
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cn
(Si
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on
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CL
li.
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>-
O
3
>
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g
U
cr
D
0) 03 *0 »0
INJN 'lAt/M aid lOHvmy
Figure lOE
The effect of added loads is shown on
the de-eyed animal (Helix lactea ) « The
relation between the velocity of a single
wave and the rate of creeping is not altered
by added loads during vertical creeping.
(See also Table Vila, page 48.)
FIG II. DIAGRAM OF THE A R R AFNGEMEINT FOR
RECORDING THE. BEHAVIOR OF THE DETACHED FOOT
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Figure 12.
A comparison of the normal (Helix lactea^)
and injected animal (adrenalin) is given in
this figure which shows the relationship
between the velocity of a single pedal \mve
and the rate of creeping.
Animal #2 Slope of line
o control 0.11
9 injected (adrenalin) 0.21
Animal #14
o control 0.29
® injected (adrenalin) 0.20
I 1 I I I I I L_
"to VS 50 55 60 6S 70 7S
FIGURE 12. VELDCITY DF PEDAL WAVE,v,
Figure 13
coTiparison of the normal and injected.
animal showing the effect of adrenalin on
the relationship between the rate of creeping
and the frequency of waves.
Animal #2 Slope of line
o = control . 0.38
« = injected (adrenalin) 0.45
Animal #14
o = control 0.22
• = injected 0.24
Figure 14
The relationship between the advance per
wave and the rate of creeping is compared with
the normal (Helix lactea) and injected animal
(adrenalin ) •
Animal #2 Slope of line
0 = control 0.17
» = injected (adrenalin) 0.18
Animal =^14
o = control 0.22
• = injected (adrenalin) 0.18
Figure 16
The relationship betv/een the velocity of a
single wave and the velocity of progression is com-
pared in the normal and s trychninized animal. This
figure shows that strychnine sulphate (1:100) in-
creases the slope of the line describing this
relationship v/hen the animal is creeping vertically
upwards and on the under surface of a horizontal
plane. It decreases the slope of this line when
creeping takes place on the upper surface of a
horizontal plane.
Plane Animal Slope of line
#6
Vertical
o
Control
Injected
( s trychnine )
0.10
0.68
Horizontal
(under surface)
o
Control
Injected
( s trychnine )
0.28
0.40
Horizontal
(upper surface)
o
Control
Injected
(strychnine )
1.22
0.87
Figure 17
A comparison of the normal and injected animal
(strycnnine sulphate 1:100) shows that strychnine
increases the slope of the line describing the
relationship betv/een the freoiiency of waves and
the velocity of creeping when cne animal is creep-
ing vertically upwards, and also when creeping on
the upper surface of a horizontal plane, r decrease
occurs when the animal is creeping on the under
surface of a horizontal plane.
Plane Animal Slope of line
#6
Vertical o = Control 0.68
» = Injected 0.88
(strychnine )
Horizontal
(under surface) o = Control 1.07
» = Injected 0.66
(strychnine)
Horizontal
(upper surface) o = Control 0.42
« = Injected 0.74
(strychnine )
i I I I I I I I I I I 1 1 1 1 LI
35 40 45 50 55 60 65 50 5.5 60 65 W 75 80 85 90 95
FIGURE 17. VELOCITY OF PROGRESSION, V "^^Vsec
Figure IS
A comparison of the normal and injected animal
(strychnine sulphate 1:100) shows that strychnine
increases the slope of the line describing the
relationships between the advance per wave and
velocity of pro!3;res3ion when the animal is creep-
ing vertically upv/ards . A slight decrease is ob-
served Y.'hen creepinging' in a horizontal plane on
the upper and lower surface.
Plane Animal Slope of line
Vertical o = Control 0.12
• = Injected 0.26
(strychnine )
Horizontal o = Control 0.32
(under surface) « = Injected 0.30
(strychnine)
Horizontal o = Control 0.22
(upper surface) « = Injected 0.14
t -
8 85 as IQO 105 lio \\5 120 \Z5 i50 05 140 i45 150 155 Q
113 116 IE3 iZ6 13.3 138 143 148 153 158 163 i&8 173 178 183 188 #
HORIZONTAL PLANE
UNDER SURFACE
115 12.0 125 130 135 14.0 145 150 155 160 #
VERTICAL PLANE
•
♦6
O = CONTROL
• = INJE.CTED /ANIMAL
1
1
1
1
]
1
1
[STRYCHNJNE SULPH/^Tt)
50 55 60 6.5 70 75 8,0 8.5 9.0 95 lOO 105
FIGURE 18. VELOCITY OF PROGRESSION, V, ""/-sec.
/
Autobiography of the Candidate
Birthplace: Cambridge, Massachusetts
Father's name: John B. Brine, born in Ca:nbridge, i.iass.
Mother's name: Mary L. Brine, born in Cambridge, Mass.
Education: Grammar School, Public School I\io.26, Kew York City
Hunter High School, New York City
A.B. 1913, Hunter College.
M.Sc. 1915, 'New York University
M.A. 1928, Radcliffe College
Positions held: Instructor in the Department of Physiology,
Hunter College, Few York City,
from 1913 to 1921.
Married: 1921, Mr. John Q. Daly, la\vyer, Boston, Mass.
/ /
//
^ ^719 02551 6941
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